Porifera: Lithistid Demospongiae (Rock Sponges)

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ISSN 1174–0043; 121

The Marine Fauna of New Zealand: Porifera: Lithistid Demospongiae (Rock Sponges) Michelle Kelly NIWA Biodiversity Memoir 121 Cover photo: Lithistid sponges on the surface of a boulder from hydrothermally active Rungapapa Knoll in the Bay of Plenty. The blue nodules and chalice-like sponges are juvenile Reidispongia coerulea (Phymatellidae), and the cream knobs are Macandrewia spinifoliata (Macandrewiidae). The boulder was retrieved using a NIWA epibenthic sled towed at 160 m depth from RV Kaharoa in 2000. P photo: NIWA NATIONAL INSTITUTE OF WATER AND ATMOSPHERIC RESEARCH (NIWA)

The Marine Fauna of New Zealand: Porifera: Lithistid Demospongiae (Rock Sponges)

Michelle Kelly

National Centre for Aquatic Biodiversity & Biosecurity National Institute of Water and Atmospheric Research (NIWA) Private Bag 109-695, Newmarket Auckland, New Zealand [email protected]

NIWA Biodiversity Memoir 121

2007 Cataloguing in Publication

KELLY, M. The marine fauna of New Zealand: Porifera: Lithistid Demospongiae (rock sponges) / by Michelle Kelly—Wellington: NIWA (National Institute of Water and Atmospheric Research), 2007 (NIWA Biodiversity Memoir, ISSN 1174-0043)

ISBN 978–0–478–23277–6

Series Editor: Dennis P. Gordon Copy edited and typeset by: Geoff Gregory, Word Therapy, Paraparaumu Printed and bound by: Graphic Press & Packaging Ltd, Levin

Received for publication—30 June 2006

 NIWA Copyright 2007 CONTENTS Page ABSTRACT...... 5 INTRODUCTION...... 6 THE NEW ZEALAND LITHISTID SPONGE FAUNA ...... …...... 7 MATERIALS AND METHODS...... ……………………………… 8 Sample collection...... ……………… 8 Spicule dimensions...... ……………………………………………… 8 Registration of type and general materials...... ……………… 9 Area of study ...... ……………………………………………… 9 Terminology...... ……………………………………………… 9 Abbreviations of institutions...... ……………………… 10 CHECKLIST OF SPECIES...... 11 SYSTEMATICS...... 12 Lithistid Demospongiae (formerly order Lithistida Schmidt, 1870)...... 12 Family Theonellidae ...... 13 Genus Discodermia...... ……………………………………… 13 Family Phymatellidae ...... ……………………………………… 15 Genus Neoaulaxinia ...... ……………………………………… 15 Genus Neosiphonia ...... ……………………………………… 22 Genus Reidispongia ...... ……………………………………… 27 Family Corallistidae...... 29 Genus Neoschrammeniella ...... ……………………………… 30 Genus Herengeria ...... ……………………………………… 34 Genus Awhiowhio n. gen. ……………………………………… ...... 39 Family Neopeltidae ...... 45 Genus Homophymia ...... ……………………………………… 45 Genus Callipelta...... ……………………………………… 47 Genus Neopelta...... ……………………………………… 50 Genus Lepidothenea...... ……………………………………… 52 Family Macandrewiidae ...... ……………………………………… 53 Genus Macandrewia ...... ……………………………………… 53 Family Pleromidae...... 54 Genus Pleroma ...... ……………………………………… 56 Family Isoraphiniidae ...... ………………………………………61 Genus Costifer ...... ……………………………………… 61 Family Scleritodermidae ...... ……………………………………… 64 Genus Microscleroderma...... ……………………………...65 Genus Scleritoderma...... 67 Genus Aciculites ...... ………………………………………69 Family Azoricidae ...... 76 Genus Leiodermatium ...... ……………………………… 77 DISCUSSION...... 88 ACKNOWLEDGMENTS...... 90 REFERENCES...... 90 APPENDIX 1. Key to diagnostic characters...... ………………………………………………………94 APPENDIX 2. Key to diagnostic characters...... ………………………………………………………95 APPENDIX 3. List of stations...... ………………………………………………………96 TAXONOMIC INDEX...... ……………………………………………… 99

Frontispiece: Lithistid sponge Aciculites pulchra Dendy (Scleritodermidae), clearly indicated by the noduled surface, on the surface of a boulder at 120 m off the southern edge of a reef off Great Barrier Island. This photograph was taken from a Remote Operated Vehicle (ROV) during the Department of Conservation’s Great Barrier Island Deep Reef Investigation in 2003. Photo: Roger Grace, Department of Conservation, Auckland Conservancy. NATIONAL INSTITUTE OF WATER AND ATMOSPHERIC RESEARCH (NIWA)

The Marine Fauna of New Zealand: Porifera: Lithistid Demospongiae (Rock Sponges)

Michelle Kelly

National Centre for Aquatic Biodiversity & Biosecurity National Institute of Water and Atmospheric Research (NIWA) Private Bag 109-695, Newmarket Auckland, New Zealand [email protected]

ABSTRACT

The lithistid Demospongiae fauna of New Zealand has been inventoried from existing and new collections, and is reviewed here and revised where necessary. Most of the 282 specimens examined were recorded from the largest collection of sponges in New Zealand, in the NIWA Invertebrate Collection, Wellington. Significant collections were also examined from the Museum of New Zealand Te Papa Tongarewa. The lithistid Demospongiae (formerly order Lithistida Schmidt, 1870) is represented in the New Zealand region by nine families, 18 genera (one of which is new to science), and 30 species (12 of which are new to science): Theonellidae (1 genus, 1 species), Phymatellidae (3 genera, 6 species), Corallistidae (3 genera, 7 species), Neopeltidae (4 genera, 4 species), Macandrewiidae (1 genus, 1 species), Pleromidae (1 genus, 3 species), Isoraphiniidae (1 genus, 1 species), Scleritodermidae (3 genera, 5 species), and Azoricidae (1 genus, 2 species). This work records the first lithistid species, Neoschrammeniella antarctica n. sp., known from polar regions, and provides the first record of the genusLeiodermatium further south than the Philippines. Two additional species of Leiodermatium described here are found only in the west-central Pacific and Micronesian deep waters, but are included for the sake of a complete review of the genus in the Pacific. New species of the previously monospecific phymatellid genera Neoaulaxinia and Neosiphonia are described, and a new corallistid genus, Awhiowhio, is recognised from New Zealand waters. All specimens were dredged from between 80 and 1700 m, but were commonest between 200 and 800 m. With the exception of one specimen from the eastern edge of the Challenger Plateau on New Zealand’s west coast, and a new species from the Ross Sea, Antarctica, all were found north of the northern edge of the Chatham Rise and in New Zealand’s northernmost waters. Known and new species are redescribed from representative New Zealand material and, in some cases, the characters used to define genera and species are redefined and clarified. In particular, ornamentation of the desma skeleton and morphology of the augmenting microscleres are emphasised for distinction at the species level.

Keywords: Porifera, lithistid Demospongiae, Lithistida, Theonellidae, Phymatellidae, Corallistidae, Neopeltidae, Macan- drewiidae, Pleromidae, Isoraphiniidae, Scleritodermidae, Azoricidae, Desmanthidae, polyphyletic group, systematics, sponges, taxonomy, new species, New Zealand

INTRODUCTION

Lithistid demosponges are a polyphyletic group com- Mayer & Hamann 2004; Piel et al. 2004). A recent survey prising 13 extant families; 36 genera are included in of the literature revealed some 40 papers reporting bio- the most recent classification (Pisera & Lévi 2002a) but logically active compounds and their synthesis from the five genera remain of uncertain status. They differ from Southwest Pacific generaAciculites , Callipelta, Discoder- other demosponges in that the dominant structural spi- mia, Microscleroderma, Neosiphonia, Pleroma, Reidispongia, cules (desmas) are articulate, forming in most species a and Scleritoderma. Of these genera, the first example of solid, rigid, heavily siliceous skeleton. These desmas are double bioalkylation of the sterol side chain at position highly diverse morphologically; the overall architecture 26 was reported from New Zealand’s Aciculites pulchra of the desma, the ornamentation of the desma surface, (Crist et al. 1983). Furthermore, Pleroma menoui Lévi & and the pattern of articulation with adjacent spicules, are Lévi, Reidispongia coerulea Lévi & Lévi, and Neosiphonia diagnostically important (Schrammen 1910, see Kelly superstes Sollas from New Caledonia were the subject of 2000a; Pisera & Lévi 2002a). considerable attention for their production of bromind- Lithistid sponges were traditionally placed in the oles (Guella et al. 1989) and antiviral (Laille et al. 1998), single order Lithistida Schmidt, 1870 owing to the com- antifungal (D’Auria et al. 1995), and cytotoxic macrolides mon possession of desmas, even though these display (D’Auria et al. 1996; Zampella et al. 1997; Carbonelli et considerable morphological diversity. For some time, al. 1999; Bassarello et al. 2000). lithistids have been recognised as polyphyletic, with Lithistid demosponges are of particular interest to several points of origin within the Demospongiae (de paleontology because many extant genera and species Laubenfels 1936; Reid 1963, 1970; Kelly-Borges & Pom- are relict from sponge faunas that were more abundant poni 1994; Pisera & Lévi 2002a). An indication of the before the present. Lithistid demosponges are known polyphyletic nature of lithistid sponges is revealed in the from the Ordovician, Silurian, Devonian, Permian, Late wide range of microscleres and ectosomal megascleres, Jurassic, Late Cretaceous, and Eocene (Pisera 2002, and desma axial geometries, which include tetraxial, 2006; Reid 2004). The Paleozoic fauna was dominated monaxial, polyaxial, and anaxial forms. by Orchocladina (extinct in the Permian) and sphaero- Familial and ordinal affiliations of lithistids with cladinid sponges (solely represented in the Recent fauna each other and with non-desma-bearing sponges are by Vetulina stalactites from the tropical Atlantic). The uncommon but can be found in the megasclere and Mesozoic (260–60 Ma) lithistid fauna was dominated by microsclere components of the skeleton (see Pisera & Tetracladina, Megamorina, and Rhizomorina, groups Lévi 2002a). The presence of triaene megascleres and that are still relatively common in Tertiary and Recent asterose microscleres in some lithistid genera clearly lithistid faunas (Pisera 2002, 2006). These sponges fos- indicates affinity with demosponge Astrophorida, and silise well because of their rigid silica skeletons; 13 this has been supported by recent DNA studies (Kelly- suborders, 34 families, and more than 200 fossil genera Borges & Pomponi 1994; Chombard et al. 1998; McIn- have been recently revised (Pisera 2002, 2006), and many erney et al. 1999). The exotylostyles of Gastrophanella more nominal genera are known. are considered to demonstrate affinity with the non- Lithistid sponges are known from almost all temper- lithistid demosponge order Hadromerida (Van Soest ate and tropical oceans but are generally restricted to & Stentoft 1988). depths greater than about 80 m in South Pacific locations A greater difficulty arises when taxa do not have including New Zealand and the Norfolk Ridge, and 150 accessory spicules, or the spicules that they have bear m in the tropical Western Atlantic region (Pomponi et only superficial resemblance to those in non-lithistid al. 2001). Apart from the proliferation of the genera Ac- demosponges. Species of Aciculites, for example, contain iculites, Neoschrammeniella, and Pleroma in the Southwest only acanthose anisostrongyles as accessory spicules, Pacific,Corallistes and Discodermia in the western tropical and Leiodermatium and Vetulina possess only desmas Atlantic, and Theonella in the Indo-Pacific, most genera and diactines, and desmas, respectively. In all three are known only from fragments or single specimens taxa, molecular evidence was required to indicate their that have been rarely, if ever, recollected (e.g. Lyidium phylogenetic relationships (Kelly-Borges & Pomponi [= Pleroma] torquilla Schmidt, 1870 from Cuba). Lithistid 1994). sponges are rarely collected from below about 1700 m The lithistid sponges are of interest to biomedical depth. science because of the great variety of pharmaceutically Prior to the present work, two major regional fau- relevant biological activities of their chemical extracts nas were known worldwide: the continental shelf and (see Pomponi 2001; Bewley et al. 1998; Munro et al. 1999; slope fauna of the tropical western Atlantic (Schmidt

1870, 1880; van Soest & Stentoft 1988; Kelly-Borges et 2005). In both locations, lithistid sponges dominate the al. 1994; Kelly-Borges & Pomponi 1994; Lehnert & van sponge fauna (Lévi 1991; Reed & Pomponi 1997; Richer Soest 1996; Pisera 1999; Pomponi et al. 2001) and the de Forges et al. 2000) between 150 and 1800 m but the seamount fauna of the Southwest Pacific including the structure and taxonomic composition of the communi- seamounts of the New Caledonian Norfolk Ridge (Lévi ties differ considerably (Pomponi et al. 2003). & Lévi 1983, 1988; Lévi 1991; Schlacher-Hoenlinger et al.

THE NEW ZEALAND LITHISTID SPONGE FAUNA

Prior to 1991, when research on the lithistid fauna of tetracladine lithistids (which are comparable to those New Zealand commenced, only two Recent lithistid of the Recent family Phymatellidae) and described a species had been described, although several more new species of Vetulina, V. oamaruensis, based upon were known (P.R. Bergquist pers. comm.). Arthur the sphaerocladine desmas. These desmas strongly Dendy described the first lithistid sponges from New resemble those found in a species of Crambe from Zealand―Aciculites pulchra Dendy, 1924 and Lepi- Spirits Bay, Northland and their conspecificity cannot dothenea incrustans (Dendy, 1924) from the vicinity of be discounted (Kelly et al. 2003). North Cape and the Three Kings Islands, respectively. Whole-body fossils of lithistid sponges are also Bergquist (1968) redescribed this material but recorded known from the mouth of the Kakanui River in the no further species. No further species were added by Oamaru district (Kelly et al. 2003). These fossils occur Dawson (1993) in his comprehensive index to the New in a volcaniclastic Ototara Limestone bed of lower Zealand Porifera. Whaingaroan age just above the top of the Mineral From 1991 onwards, many of the lithistid species Breccia Member of the Deborah Volcanics (31.6 Ma) described by Lévi and Lévi (1983, 1988) from the south (Dickey 1968; Daesch et al. 1970). The body fossils were New Caledonian slope and seamounts were progres- found to be very similar morphologically to the living sively discovered in the New Zealand region (Kelly et pleromid sponge Pleroma aotea Kelly when compared al. 1999; Kelly 2000b; Kelly 2001a,b; Pomponi et al. 2001; morphometrically to known extant sponges from New Kelly 2003; Kelly et al. 2003; Kelly & Buckeridge 2003; Zealand and New Caledonia. Kelly et al. (2003) consid- Kelly 2004; Kelly & Tubbs 2006; Kelly et al. in press), ered them to be conspecific with living sponges from along with several undescribed species. A voyage to deepwater seamounts and banks off northeastern New the seamount region south of New Caledonia in Au- Zealand, citing several additional examples from the gust 1999, courtesy of IRD, Nouméa, assisted greatly substantial record of lithistid sponges in the Oamaru in the author’s understanding of the faunal relation- Diatomite. ships between the two countries and of lithistid sponge Lithistid microfossil spicules and partial body ecology in general. fossils are also known from the Tutuiri Greensand Homophymia stipitata Kelly, 2000a, discovered in (Teurian–basal Waipawan) in the Chatham Islands New Zealand waters, was only the second known spe- (Buckeridge in Campbell et al. 1993; Buckeridge & cies of the genus, which was known previously only Kelly 2002; Kelly & Buckeridge 2005). The spicules from a single species from Madagascar and Réunion. are trapped within the skeletons of fossil hexactinel- Also recently recognised in national collections was lid sponges. These are remarkable for their diversity, Pleroma aotea Kelly, 2003a, the third species to be and the combination of lithistid, astrophorid, and described in a genus principally known from New hexactinellid taxa indicates a paleoenvironment very Caledonian waters. similar to that found today at depths of 500–800 m on Prior to the work of Dendy (1924), Hinde and the Chatham Rise (Buckeridge & Kelly 2005) and in the Holmes (1892) described a species-rich assemblage tropical Atlantic (cf. Pomponi et al. 2001). based on siliceous spicules from marine diatomaceous Reasonably well-preserved sponge body fossils sediments now known to be from the early Runangan resembling lithistid Corallistidae, Isoraphiniidae, (Late Eocene) horizon within the Oamaru Diatomite Pleromidae, Phymatellidae, and Scleritodermidae, Member of the basaltic Waireka Volcanics (c. 35 Ma) at have also been found in the Red Bluff Tuff (Teur- Oamaru in North Otago (Suggate et al. 1978; Edwards ian–Waipawan) on Chatham Island (Buckeridge in 1991). Amongst the many non-lithistids represented Campbell et al. 1993; Buckeridge & Kelly 2006), provid- were microfossil spicules of what were considered ing an interesting record for this group that straddles to be a species of Lyidium [= Pleroma], and species of the Mesozoic–Cenozoic boundary (Maastrichtian to Corallistes, Discodermia, and Theonella were illustrated. early Ypresian) (Kelly et al. 2006). Hinde and Holmes (1892) also illustrated desmas of

MATERIALS AND METHODS

SAMPLE COLLECTION Specimens were either frozen immediately upon collection or preserved in 50% isopropanol and pre- Sponges were collected by rock dredge from north- pared for histology and scanning electron microscopy eastern New Zealand waters by the National Institute as in Kelly-Borges and Vacelet (1995). Specimens are of Water & Atmospheric Research (NIWA) research stored for the long term in 70% ethanol. vessels Tangaroa (1967–1989) and Kaharoa (1998) (Fig. 1). Specimens of known New Zealand species collected by the author on the IRD cruise LITHIST (RV Alis, SPICULE DIMENSIONS August 1999) on the New Caledonian Norfolk Ridge Whole desmas are notoriously difficult to measure seamounts have been included in the study insofar as because of their intimate articulation with adjacent they represent a natural northern extension of the range spicules and their frequent irregular morphology. The of the New Zealand species. NIWA, NMNZ, and IRD size of the desma is expressed here as the approximate station data are listed in Appendix 3; only the station size of the overall spicule, and as the mean clone length name is listed in the sections describing material exam- and thickness. All spicule measurements are expressed ined. Data for other stations are listed in the text. in µm. Microsclere dimensions recorded in this work

Figure 1. Locality maps, showing the collection stations for rock sponges in New Zealand and Antarctic waters.

are generally given as mean length (range of length TERMINOLOGY measurements) and mean width (range of width measurements) using the measurements of 10–15 spicules Specialist terminology for lithistid sponges follows where possible. In the case of the diactinal megascleres, Kelly (2000a) and Pisera and Lévi (2002a). General it was very difficult to measure whole spicules as they terminology for sponges is available in Boury-Esnault were typically fragmented. Dimensions are listed in and Rützler (1997), but some terms are included here tabular format in the description of each species. The for convenience. colour of the sponges in life and in ethanol is given by acantho—prefix meaning roughened or microspined, name and also code according to the Reinhold Colour e.g. acanthoxea, acanthorhabd Atlas (Kornerup & Wanscher 1961). amphiaster—microsclere with equal numbers of rays projecting from both ends of an elongate centrum, equidistant from the centre; the prefix defines the REGISTRATION OF TYPE AND GENERAL shape of the ray, whether fine and pointed (oxy-) MATERIALS or rounded and robust (strongylo-) Primary and secondary type materials of new species, anaxial desma—devoid of a crepis (see sphaeroclone) and additional material, are deposited in the NIWA clad(ome)—see triaene Invertebrate Collection at the National Institute of clone—ray-like arm of a desma that is partly (crepis Water & Atmospheric Research (NIWA; formerly extends a short way along the clone from the proxi- New Zealand Oceanographic Institute/ NIWA), Greta mal end) or entirely anaxial (devoid of a crepis); the Point, Wellington. Previously published type and gen- number of clones is determined by the desma geom- eral registration numbers for the species Homophymia etry (modified from Pisera & Lévi 2002a) stipitata and Pleroma aotea are retained in this work, but crepis—the inceptional body or axial filament of the their new registration numbers (NIWA----) are given desma that is visible as a short thread-like canal in in parentheses for reference. Some type and additional monocrepid desmas, or cruciform canals in tetracrepid material collected by the Coral Reef Research Founda- desmas (modified from Pisera & Lévi 2002a) tion was previously registered at the Natural History desma—articulate choanosomal megasclere with a Museum, London (BMNH) and the practice is con- variety of geometries, often with complex inter- tinued here if the material is from non-New Zealand connected morphology, often highly ornamented; locations. Some material was previously registered in found in lithistid sponges (modified from Pisera & the Museum of New Zealand Te Papa Tongarewa (for- Lévi 2002a) merly National Museum of New Zealand, NMNZ) and dichotriaene—an ectosomal spicule of triaenose sym- retains the prefix NMNZ Por. ---. The prefix IRDR---- is metry (see triaene) with a regular cladome of three for material sourced with permission from Institut de branches (protoclads) that can divide further to Recherche pour le Développment (IRD) Noumea, col- form usually two clads (deuteroclads); the clads lections. Registration numbers are cited in the text. can be smooth, tuberculate, or spinose and are tangential to the sponge surface; the rhad can be short AREA OF STUDY or long and is always perpendicular to the clads or cladome, penetrating the sponge surface; axial Sponge specimens in this monograph are from col- canals extend the whole length of the rhabd and lections made by NIWA (formerly as New Zealand clads (modified from Pisera & Lévi 2002a) Oceanographic Institute of the DSIR) and the National dicranoclone—see monocrepid desma; arch-shaped and Museum of New Zealand Te Papa Tongarewa. The col- bearing well-developed fungiform tubercles, can lection area (Fig. 1) extends from 24° to 74° S and 155° be bi-, tri-, or sometimes tetrapodial (four-footed); E to 178° W, covering essentially the Lord Howe Rise, zygomes are terminal on clones and articulate with Dampier Ridge, South Fiji Basin, parts of the Norfolk the upper tubercles of clones from adjacent desmas, Ridge, the Southwest Pacific Basin, Chatham Rise e.g. in Corallistes and Herengeria; (Corallistidae) and Subantarctic Slope in the south, and the Ross Sea, (modified from Pisera & Lévi 2002a) Antarctica (Appendix 3). The region includes Norfolk discotriaene—ectosomal megasclere with tangential Island, Lord Howe Island, and the Kermadec Islands cladome forming a flat or slightly concave oval disc, in the north. Depths range from 80 to nearly 1680 m. In margins can be even or incised; rhabd is short, crepis addition to these New Zealand collections, specimens is tetraxial, e.g. in Discodermia (Theonellidae) (modi- collected by the author from the New Caledonian fied from Pisera & Lévi 2002a) Norfolk Ridge seamounts, the Lord Howe Seamount ectosome—refers to the region just below and at the Chain, and Palau, Micronesia, were included in the surface of the sponge; frequently referred to in monograph to complete the present revision. terms of the architecture of this region as it is often diagnostic and quite different from the under

lying choanosome (see Boury-Esnault & Rützler structural skeletal component, the mesosclere― 1997) a medium-sized spicule found in Homosclero- heloclone—see monocrepid desma; elongate, oxea-like, phorida, and microsclere―the smallest size of spicule smooth with notch-shaped zygomes, e.g. in Costifer (see Boury-Esnault & Rützler 1997) (Isoraphiniidae) streptaster—microsclere with rays projecting irregu- megaclone—see monocrepid desma; arched and smooth larly along an elongate centrum, the number of rays with cup- or saddle-shaped zygomes, e.g. in Pleroma ranging from numerous to only a few (Pleromidae) tetraclone—tetracrepid desma; may be smooth or tu- megasclere—see spicule berculate; very regular, often with four clones, but microsclere—see spicule often secondarily modified monaxial desma—see monocrepid desma tetracrepid desma—tetraxial desma, tetraclone; the monocrepid desma—monaxial desma; the monocrepidial tetracrepidial nature is revealed by the crepis which nature is revealed by the crepis which is a short is cruciform; zygomes mostly terminal on clones straight canal in the middle of the clone (modified (modified from Pisera & Lévi 2002a) from Pisera & Lévi 2002a) (see megaclone, heloclone, tetraxial desma—see tetracrepid desma dicranoclone, rhizoclone) triaene—general term for a tetractinal megasclere with phyllotriaene—ectosomal spicule with a single ray four rays (clads) emanating from a single point in called a rhabd, usually perpendicular to the sponge three axial planes (cladome), having one unequal surface and penetrating it, and three others are more ray (rhabd, rhabdome) that is commonly much or less flat and tangential to the sponge surface, longer than the other three rays (see Boury-Esnault termed the cladome; the cladome is branched in an & Rützler 1997) irregular manner with the clads resembling feathers; tuberculate—warty; tubercles have a rounded apex the crepis is tetraxial and very short in the radiat- and straight or restricted sides, forming a fungiform ing branches of the cladome (modified from Pisera (mushroom-shaped) tubercle & Lévi 2002a) zygome—the articulating clasp formed between the pseudodiscotriaene—ectosomal megasclere resem- tips of clones of adjacent desmas (modified from bling a discotriaene and analogous to it, but is Pisera & Lévi 2002a) monocrepid with the crepis located in the rhabd or cladome (e.g. Neopeltidae) (modified from Pisera ABBREVIATIONS OF INSTITUTIONS & Lévi 2002a) pseudophyllotriaene—ectosomal spicule closely re- AUT Auckland University of Technology. sembling a phyllotriaene and analogous to it, but CRRF Coral Reef Research Foundation, based in monocrepid (crepis may be located in the rhabd the Republic of Palau, Micronesia. or in the cladome) (modified from Pisera & Lévi IRD Institut de Recherche pour le Développe- 2002a) ment (formerly ORSTOM). pseudospheraster—microscleres with slightly acentri- MNHN Muséum National d’Histoire Naturelle, cally projecting massive spiny rays and a swollen Paris. centrum, that resemble spherasters, but are most NHM The Natural History Museum (formerly probably modified amphiasters British Museum (Natural History)), London. —monocrepid desma; complex shape pseudotetraclone NIWA National Institute of Water and Atmos- with elongate clones resembling a tetraclone or pheric Research (including the former New rhizoclone; the term megarhizoclone has been used Zealand Oceanographic Institute, NIWA), to describe a variety of non-related desmas that superficially resemble rhizoclones; Pisera and Lévi Greta Point, Wellington. (2002a) recommended that this term be abandoned NMNZ Museum of New Zealand Te Papa Ton- and favoured the use of pseudotetraclone instead garewa (formerly National Museum of of megarhizoclone (see Kelly 2000) New Zealand), Wellington. rhab(dome)—see triaene 0CDN Sample numbers for material collected by rhizoclone—monocrepid desma; usually with numerous the Coral Reef Research Foundation for spines and or tubercles that serve as lateral zygomes United States National Cancer Institute (modified from Pisera & Lévi 2002a) shallow-water collection programme. A sphaeroclone—anaxial desma; in which several ray-like complete collection of all 0CDN sponge arms extend from a globular centre that may be specimens is located at the Smithsonian spinose (modified from Pisera & Lévi 2002a) Institution (U.S. National Museum), and spicule—opaline-silica or calcium carbonate com- with the author. ponent of the poriferan skeleton; often of elabo- USNM U.S. National Museum (Smithsonian Insti- rate morphology and design; three general size tution). categories include the megasclere―the primary

10 CHECKLIST OF SPECIES

Class DEMOSPONGIAE Family PLEROMIDAE Family AZORICIDAE Pleroma aotea Kelly Leiodermatium colini n. sp.* Pleroma menoui Lévi & Lévi Leiodermatium dampieri n. sp. Pleroma turbinatum Sollas Leiodermatium intermedia (Sollas, 1888)* Leiodermatium linea n. sp. Family PHYMATELLIDAE Neoaulaxinia clavata (Lévi & Lévi) Family CORALLISTIDAE Neoaulaxinia persicum n. sp. Awhiowhio osheai n. gen. n. sp. Neoaulaxinia zingiberadix n. sp. Awhiowhio sepulchrum n. gen. n. sp. Neosiphonia motukawanui n. sp. Awhiowhio unda n. gen. n. sp. Neosiphonia superstes Sollas Herengeria auriculata Lévi & Lévi Reidispongia coerulea Lévi & Lévi Herengeria vasiformis Schlacher-Hoenlinger et al. Neoschrammeniella antarctica n. sp. Family Scleritodermidae Neoschrammeniella fulvodesmus (Lévi & Lévi) Aciculites manawatawhi n. sp. Aciculites pulchra Dendy Family ISORAPHINIIDAE Aciculites sulcus n. sp. Costifer wilsoni Lévi Microscleroderma novaezelandiae n. sp. Scleritoderma flabelliformis Sollas Family MACANDREWIIDAE Family THEONELLIDAE Macandrewia spinifoliata Lévi & Lévi Discodermia proliferans Lévi & Lévi Family NEOPELTIDAE Callipelta punctata Lévi & Lévi Homophymia stipitata Kelly Lepidothenea incrustans (Dendy) Incertae sedis Neopelta pulvinus n. sp.

* The two asterisked species of Leiodermatium described in this monograph are found only in the west-central Pacific and Micronesian deep waters, but are included in here for the sake of a complete review of the genus in the Pacific.

11 SYSTEMATICS

The lithistid Demospongiae have recently been revised Family THEONELLIDAE Lendenfeld, 1903 by Pisera (2002) and Pisera and Lévi (2002a-o) and are considered by them to comprise 13 extant families with Theonellidae Lendenfeld, 1903: 125 (in part); Lendenfeld 1907: 343 (part); Wilson 1925: 447; Lévi 1991: 79; Pisera 26 valid genera. The systematics scheme that is used in & Lévi 2002e: 327. this volume follows these recent revisions. The reader Discoderminae Schrammen, 1910: 97. is referred to the Systema Porifera―a major publication Discodermiidae: Schrammen 1924: 37, 48; de Laubenfels for full family and genus-level synonymies. 1955: E58 (part). Lithistid taxa are extremely difficult to identify without practice. To assist field and laboratory iden- Polymorphic; choanosomal spicules are tetraclone tification, a summary section termed Key Diagnostic desmas that, in several genera, may be non-articulated Characters is given immediately after the Remarks with neighbouring desmas, being tetralophose in following the formal description of each species. The general shape; ectosomal spicules are phyllotriaenes, characters listed are those that can be used rapidly in discotriaenes, or variations between these forms; cho- succession in the field and laboratory to determine ge- anosomal diactinal megascleres often have a strongy- nus and species in relation to morphologically similar lote morphology, some with tylote (hammer-like) other species. Keys to the diagnostic field and micro- proximal and blunt stylote distal ends, or these may be scopic characters of New Zealand lithistid sponges is long thin oxeas; microscleres include acanthorhabds, given in Appendices 1 and 2. acanthostrongyles, microxeas, streptasters, or pseu- dospherasters (modified from Pisera & Lévi 2002e). Lithistid Demospongiae (formerly order Lithistida Schmidt, 1870) Discodermia du Bocage, 1870

Polyphyletic group of encrusting, pedunculate, fungi- Discodermia du Bocage, 1870: 1; Zittel 1878: 37, 87; Sollas 1888: form, auricular (ear-shaped), fan-shaped, cup-shaped, 292 (part); Pisera & Lévi 2002e. or massive-amorphous demosponges characterised by the presence of articulated desma megascleres of Theonellidae with ectosomal discotriaenes; regular diverse morphology that render most such sponges tetraclone desmas with smooth or tuberculate clones rigid and rock-like. Choanosomal desmas are morpho- and zygoses; choanosomal megascleres long oxeote or logically diverse and comprise tetraxial (tetraclone), stylote; microscleres are large curved acanthoxeas and monaxial (rhizoclone, megaclone, dicranoclone, helo- smaller acanthorhabds. clone, or various complex branched forms), polyaxial, or anaxial (sphaeroclone) geometry. Additional Remarks: Hinde and Holmes (1892) illustrated several choanosomal megascleres may include long hair-like microfossil discotriaenes from the Oamaru Diatomite oxeas. Ectosomal spicules may include dichotriaenes, (c. 35 Ma) at Oamaru, North Otago, indicating that phyllotriaenes, pseudophyllotriaenes, discotriaenes, the genus Discodermia (or Theonella as per Hinde & pseudodiscotriaenes, and various roughened oxea- Holmes, 1892, pl. 14, figs 4, 5, 8–11) was present in the like spicules. Microscleres include a combination of southern New Zealand location that is now Oamaru, spirasters, streptasters, amphiasters, sigmaspires, during the late Eocene. One microfossil was attributed roughened oxeas or raphides (after Pisera & Lévi to Discodermia sinuosa Carter, 1881, originally described 2002a). from the Gulf of Manaar and the southeast coast of Sri Lanka (Hinde & Holmes 1892, pl. 14, fig. 12). The discotriaene of this species also has indented margins, but Remarks: The informal group lithistid Demospongiae has recently been revised by Pisera (2002) and Pisera the similarity of the illustration to the phyllotriaenes and Lévi (2002a-o) and is considered by them to contain of Macandrewia spinifoliata Lévi & Lévi cannot be over- 13 extant families with 26 valid genera (five of uncer- looked. Conspecificity with an extant species from the tain affinity). The classification scheme that is used in Western Indian Ocean is less likely. this volume, and the sequence of families and genera, follows these recent revisions. For a full discussion of Type species: Dactylocalyx polydiscus Bowerbank, 1869. each family and genus see Pisera (2002) and Pisera and Lévi (2002a-o).

12 Discodermia proliferans Lévi & Lévi, 1983 perpendicular to the surface but do not pierce the (Fig. 2) surface. Megascleres (Table 1) robust tetraclone desmas (Fig. Discodermia proliferans: Lévi & Lévi, 1983: 121, pl. 6 (fig. 4), 2B–D) with thick short smooth tubercles covering the pl. 7 (figs 1, 2, 4–9); Schlacher-Hoenlinger et al. 2005: 681, entire spicule, zygoses are very tight and intricate. figs 4A, 17, 31. Discotriaenes (Fig. 2C–E) with the plane of the disc uneven and the edges frequently faintly crenulate, or Material examined: deeply incised owing to obstruction from the rhabds of NIWA Stations: S572 (NIWA18212); X768 adjacent discotriaenes, rhabds short and conical. Diac- (NIWA18213). tinal megascleres with strongylote proximal and distal IRD Stations: LITHIST Cruise (August 1999) DW5 termini, with spicule curved and abruptly slimming at (NIWA18480), DW7 [NIWA18481, NIWA18482 the ends and curveing inwards, uncommon. (IRDR1829)]. Microscleres (Table 1) large acanthoxeas, faintly centrotylote, and occasionally sharply angled around Distribution: Passe de la Havannah (holotype), Introu- midsection of spicule (Fig. 2C). Acanthorhabds with vable, Stylaster, and Éponge Seamounts, south New strongylote ends, occasionally centrotylote (Fig. 2F). Caledonian slope of Norfolk Ridge; west of Three Kings Ridge, northern New Zealand; Hikurangi Plateau, east Remarks: All specimens examined conform to the holo- of East Cape, New Zealand. type of Discodermia proliferans from New Caledonia (Lévi & Lévi 1983) and the species is typical of the genus Habitat: Sponges were dredged from hard surfaces with distinct discotriaenes and two forms of acanthose on the tops of seamounts and continental margins; the microsclere. However, although some specimens from substratum of sponges from NIWA Stn X768 was grey Éponge Seamount DW7 (NIWA18481–2) had the same mudstone. Depth range 175–936 m. morphology, they had abundant diactinal spicules and differred in the possession of smaller thinner acanthox- Description: eas and acanthorhabds, and the desmas were smaller Morphology arborescent, often a curved arched column and ridged rather than distinctly tuberculate. with lateral tuberculate projections, frequently joined Tuberculate ornamentation on the clones of Atlan- to neighbouring sponges (Fig. 2A). tic and Pacific species of Discodermia spp. is rare. The Dimensions 40–65 mm long, column 5–10 mm thick, only other species of Discodermia known to the author 20 mm thick at widest point including projections. that has annular rings or partial ridges and slight Texture stony, granular. nodules on the clones is an undescribed species from Surface smooth with subdermal aquiferous canals Jamaica (Kelly & Pomponi unpubl.). NIWA18481 and visible. NIWA18482 from Éponge Seamount (DW7) are very Colour cinnamon brown in life (6D6), ivory in close to Discodermia gorgonoides Burton, 1928 described ethanol (4B3). from the Andaman Islands in the Indian Ocean, except Choanosomal skeleton is composed of a dense articu- that the latter species lacks acanthoxeas. The holotype lation of robust tetraclone desmas interspersed with of D. gorgonoides was only a fragment of what was dense masses of large acanthorhabds (Fig. 2B,C). thought to be a branching sponge with oxeas and Ectosomal skeleton is composed of discotriaenes, notched discotriaenes, but these are only about 180 acanthoxeas, and acanthorhabds (Fig. 2D). Occasional µm wide. The nodules on D. gorgonoides are not as diactinal spicules traverse the deeper choanosome pronounced as on typical specimens of D. proliferans.

Table 1. Spicule dimensions (µm) of Discodermia proliferans Lévi & Lévi, 1983. Desmas Discotriaenes Diactines Acanthoxeas Acanthorhabds (Clone L/Thickness) (Max disc ø) (L) (L/Thickness) (L/Thickness) New Zealand NIWA18212 300 (250–400)/ 243 (190–243) 400–1500 104 (95–112)/3–4 13 (10–15)/3–4 100 (80–150) NIWA18213 340 (250–400)/100 540 (375–725) 400–1500 90 (90–100)/5–6 8 (7–10)/3–4 New Caledonia NIWA18480 300(200-400)/100 420 (240–620) 530 86 (70–110)/2–3 10 (7–13)/2–3 Lévi & Lévi (1983) 300/50 200–300, 250–400 — 60–80/3–5 8–11 S.-H. et al. (2005) 500–600/80–90 492–673 — 84–117/6–7 10–14/5–6

13 Figure 2. Discodermia proliferans Lévi & Lévi, 1983.

14 Branching morphology in species of Discodermia is two new species of Neoaulaxinia, and one of Neosiphona not uncommon although most species are cup-shaped has extended the list of diagnostic characters that en- or hemispherical with massive or nodular morphology. able easier differentiation at the generic and species Discodermia vermicularis Döderlein, 1884 from Japan has level within the family. digitate outgrowths arising from an encrusting base It is interesting to note that microfossil tetraclones and D. ramifera Topsent, 1892 from the Azores is digi- and ectosomal triaenes indistinguishable from those of tate. One of the commonest species of Discodermia in extant phymatellid genera were found in the Tutuiri the central western tropical Atlantic is digitate (Kelly- Greensand (Teurian–basal Waipawan) in the Chatham Borgest et al. 1994) and a further (undescribed) tiny Islands (Buckeridge & Kelly 2005, fig. 4C), and illus- branching species is known from the Gulf of Mexico trated from the Oamaru Diatomite at Oamaru in North (Kelly & Pomponi unpubl.). Otago (Hinde & Holmes 1892; pl. 13, figs 28–30), con- Specimens from New Zealand and New Caledonia firming a more southern distribution for Phymatellidae (Lévi & Lévi 1983; Schlacher-Hoenlinger et al. 2005) during the Paleogene. present a broad range of spicule dimensions, particularly the desmas, discotriaenes, and acanthoxeas, but Neoaulaxinia Pisera & Lévi, 2002 are considered to be conspecific despite this. Spherical Phymatellidae with a restricted base, elongated to various degrees, forming clavate (club- Key diagnostic characters: shaped), multiclavate, tuberose, or apple-shaped spe- • Sponge forms a small nodulose branch or thick ir- cies; subdermal aquiferous canals visible beneath the regular nodulose lump. ectosome converge on a single membranous oscule in • Surface of sponge covered with disc-like triaenes. a shallow apical depression; desmas are simple tetra- • Desmas are heavily tuberculate. clones with smooth clones that vary in the degree of • One form of small microrhabd microsclere. ornamentation (single to multiple fungiform tubercles, to spires of tubercles). Desmas branch at the terminus Family PHYMATELLIDAE Schrammen, 1910 of the clones. Zygosis is highly developed in mature spicules, forming an enlarged complex tangle; zygosis Phymatellinae Schrammen, 1910: 72. is positioned terminally and laterally with saddle for- Phymatellidae: Schrammen 1924; Lévi 1991: 79; Pisera & mations; ectosomal dichotriaenes with finely tapered Lévi 2002k: 366. frequently irregularly curved clades and rhabd; three clearly differentiated microscleres, a robust strongy- Clavate, multiclavate (tuberose or corm-like), spheri- loamphiaster restricted to an ectosomal crust, a large cal with a restricted base, and cup-shaped to foliose streptaster with only a few conical rays and a smaller sponges; desmas tetraclones with large, smooth, tuber- streptaster with numerous fine rays, restricted to the culate or semi-spinose clones; in some genera clones choanosome. are branched and ramified; ectosomal megascleres are relatively short-shafted dichotriaenes and long thin Type species: Aulaxinia clavata Lévi & Lévi, 1988. oxeas; ectosomal microscleres are acanthose strongyloamphiasters with robust rays, some with streptaster or spiraster modifications, choanosomal microscleres Neoaulaxinia clavata (Lévi & Lévi, 1988) are acanthose amphiasters that merge into streptasters (Fig. 3) with slender oxeote rays, often in two size categories. Aulaxinia clavata Lévi & Lévi, 1988: 243; Pisera & Lévi 2002k: 366, figs 1–5. Remarks: The Phymatellidae contains three valid extant genera differentiated primarily by their habit Material examined: (Pisera & Lévi 2002k), described as being clavate or NIWA Stations: KAH0204/07 (NIWA18214); apple-shaped in Neoaulaxinia Pisera & Lévi, spherical KAH0204/47 (NIWA25644). with a stem in Neosiphonia Sollas, and cup-shaped or IRD Stations: LITHIST Cruise (August 1999) CP8 lamellate in Reidispongia Lévi & Lévi. New specimens (NIWA18483), CP9 (NIWA18484). of Neoaulaxinia clavata (Lévi & Lévi), Neosiphonia superstes Sollas, and Reidispongia coerulea Lévi & Lévi Distribution: Station DW66 (BIOCAL) (holotype), and have enabled clarification of the skeletal differences Éponge Seamount, south New Caledonian slope of between the genera, focusing on the degree of and Norfolk Ridge; Seamount 441 and Cavalli Seamount, nature of ornamentation of the tetraclone desmas and northeastern New Zealand. the definition of the microscleres. The recognition of

15 Figure 3. Neoaulaxinia clavata (Lévi & Lévi, 1988).

16 Habitat: Dredged from hard coral substratum on the streptasters in two distinct size categories with fine tops of seamounts. Depth range 505–880 m. oxeote rays (Table 2, Ms 2.1, 2.2, Fig. 3F,G). NIWA18483 has spirasters with 2–4 whorls.

Description: Remarks: Only a single specimen has been recorded Morphology club-shaped, slightly thicker in the upper from New Zealand waters; the description and illustra- third, tapering to a rounded apex (Fig. 3A), or multicla- tions provided are based on this and specimens col- vate in older specimens, with a rounded stem, apple- lected from New Caledonian seamounts. The morphol- shaped apical bulb, and lateral club-shaped protuber- ogy and skeletal details of the New Zealand specimen ances (Fig. 3B). A single small membranous oscule is are very close to the New Caledonian holotype except visible on the apex of each specimen (or protuberance), that the cladome of the dichotriaenes is frequently less than 1 mm diameter; apical cavity visible only in malformed with rounded strongylote clades (Fig. 3F) some specimens, and then only 1–2 mm deep. and the cladome is occasionally monotrianeose, with Dimensions of single clubs 60–70 mm long, 15–20 unbranched clades. Excellent illustrations of the details mm thick at broadest; multiclavate specimen 160 mm of this species in New Caledonia were provided by Lévi high and 80 mm wide. and Lévi (1988) and Pisera and Lévi (2002k). Texture firm, slightly compressible. A large multiclavate specimen was recovered from Surface smooth, glistening, with longitudinal sub- Éponge Seamount on the New Caledonian end of the dermal aquiferous canals clearly visible. Norfolk Ridge that is in every aspect of the skeletal Colour in life and in ethanol ivory (4B3). and spicule morphology and dimensions, identical to Choanosomal skeleton relatively cavernous with aquif- described and new material (Fig. 3B). The size of this erous canals permeating the skeleton, and composed of specimen indicates that it might be older than the sin- weakly articulated dispersed desmas, between which gle club-shaped specimens typically recovered in the are abundant microscleres of the largest category. past, and by this we have an indication of the mature Ectosomal skeleton encrusted with robust microscle- morphology. res, interspersed with the cladomes of the dichotriaenes. Large robust oxeas arise in tracts from within the Key diagnostic characters: deeper choanosome, but do not pierce the surface. Megascleres (Table 2) tetraclone desmas with smooth • Sponge typically forms an elongate club with a even clones that branch just before they terminate, each single oscule at the apex. branch dividing into multiple tubercular fingers (Fig. • Desmas branch terminally and laterally; zygomes 3C); tubercles are typically single and slightly fungi- are lightly tuberculate. form (Fig. 3D), zygosis with adjacent spicules is weak, • There may be spirasters amongst the micro- and can be positioned laterally. Dichotriaenes (Fig. 3E) scleres. have finely pointed clads and are relatively thin overall. The individual deuteroclads are curved and separated Neoaulaxinia zingiberadix n. sp. (Fig. 4) by approximately 45°. In a New Zealand specimen, modification of the cladome to strongylote ends was Material examined: relatively common (Fig. 3F). Diactinal spicules are relatively thick, finely pointed oxeas (Fig. 3C,E). Type material: Holotype BMNH 1995.3.30.3 from Microscleres (ectosomal) are strongyloamphiasters NIWA Stn J953. (Table 2, Ms 1, Fig. 3G), choanosomal microscleres are

Table 2. Spicule dimensions (µm) of Neoaulaxinia clavata (Lévi & Lévi, 1988). Desma (Clone L/T) Cladome (Max ø) Diactines Ms 1 (L) Ms 2.1 (L) Spirasters Desma (Overall size) Rhabd (L) (L) Ms 2.2 (L) (L) New Zealand NIWA18214 375–550/50–60 430 (325–625) — 14 (10–20) 36 (30–40) — 575–750 320 (250–375) 20 (15–25) New Caledonia NIWA18483 200–400/40–60 478 (375–575) 350-4000 14 (10–18) 42 (38–48) 15 (8–18) 575–900 overall 423 (325–475) 15 (13–18) Lévi & Lévi (1988) 300–600/100 250–600 700–4000 13–15 20–40 — 600–1000 overall 200–480

17 Figure 4. Neoaulaxinia zingiberadix n. sp.

18 Type locality: Western continental slope, Northland, Etymology: Named for the morphology of the sponge New Zealand. that resembles the root (Latin, radix) of the ginger plant (Latin, zingiber). Distribution: Western continental slope, Northland, New Zealand. Remarks: Neoaulaxinia zingiberadix is only the second Habitat: Dredged from hard coral substratum on the species described in this genus, known only from tops of seamounts. Depth range 260–270 m. the Southwest Pacific. The primary character that differentiates this new species of Neoaulaxinia is the Description: morphology of the sponge. Although only a single Morphology tuberose or corm-like with multiple pro- specimen has been recovered, it is sufficiently distinct tuberances of irregular size and shape emerging from from multiclavate specimens of N. clavata to warrant the mass of the sponge; the point of attachment is a the establishment of a new species. short stem (Fig. 4A); multiple membranous oscules Neoaulaxinia zingiberadix is much less regularly are visible on apex of each protuberance. No apical clavate than N. clavata and the protuberances vary cavity around oscules. greatly in size and shape (Fig. 4A). Spicule morphology Dimensions 50 mm high, 30 mm thick. and dimensions also differ considerably; the strongy- Texture firm, almost stony. loamphiasters are larger in the new species than in Surface smooth. N. clavata, but the larger category of streptasters in N. Colour in life unknown, dried, cream (4A2). zingiberadix is generally smaller than N. clavata. The Choanosomal skeleton relatively cavernous and is dichotriaenes differ dramatically, in the smaller length composed of relatively firmly articulated desmas (Fig. of the rhabd in N. zingiberadix and in the general shape 4B). and orientation of the deuteroclads. In N. zingiberadix Ectosomal skeleton encrusted with microscleres. Ox- the individual deuteroclads are relatively straight and eas arise in tracts from within the deeper choanosome, separated by a greater than 45° angle, so that the clads but do not pierce the surface. lie close together (Fig. 4C). In N. clavata the individual Megascleres (Table 3) tetraclone desmas with smooth deuteroclads are curved and emerge with an approxi- even clones that branch just before they terminate, each mately 45° angle, so that the clads are well separated. branch dividing into multiple fingers; tubercles are As for N. clavata, the desmas are sparsely tuberculate fungiform and frequently multiple (Fig. 4B). Zygosis and the tubercles are usually fungiform, but in N. zin- is strong and situated laterally and terminally. Dicho- giberadix the tubercles are often double or triple. The triaenes have a short rhabd (Fig. 4B,C). The individual articulation of these desmas is very intricate and the deuteroclads are relatively straight and separated by sponge is a lot harder as a result. The morphology of a greater than 45° angle, so that the individual clads the terminal zygosis in the new species is that of sliding lie close together (Fig. 4C). Diactinal spicules are rela- clasped fingers (zygosis is lateral as well as terminal) tively thick centrally, but taper abruptly to strongylote and the lateral zygoses are saddle shaped. ends. Microscleres (choanosomal) are amphiasters with Key diagnostic characters: streptaster modifications in two close size categories, differentiated primarily on the number of rays on the • Sponge forms a tuberose or corm-like mass. spicule; the smaller category has more rays than the • Desma clones branch terminally and laterally and larger category (Table 3, Ms 2.1, 2.2) (Fig. 4D). Ectos- are lightly tuberculate, tubercles are frequently omal microscleres are strongyloamphiasters with a few double. robust rays (Table 3, Ms 1, Fig. 4E). • Dichotriaenes have a short rhab and clads emerge from the protorhabd well separated.

Table 3. Spicule dimensions (µm) of Neoaulaxinia zingiberadix n. sp. Desma (Clone L/T) Cladome (Max ø) Diactines Ms 1 (L) Ms 2.1 (L) Desma (Overall size) Rhabd (L) (L) Ms 2.2 (L)

New Zealand BMNH 1995.3.30.3 300–500/50–70 490 (250–775) 775+ 19 (18–23) 29 (23–35) (holotype) 500–875 158 (100–300) 18 (15–20)

19 Figure 5. Neoaulaxinia persicum n. sp.

20 Neoaulaxinia persicum n. sp. (Fig. 5) grew. Depth range 503–1680 m.

Description: Material examined: Morphology spherical (Fig. 5A) to egg-shaped (Fig. Type material: Holotype NIWA18215 from NIWA 5B,C); the base is a thin skirt, or encompassing branches Stn X140; paratypes NIWA18221 from NIWA of scleractinian corals. A single membranous oscule Stn KAH0204/29; NIWA18222 from NIWA Stn is situated at the apex of the sponge surrounded by KAH0204/30; NIWA18216 from NIWA Stn deep subdermal aquiferous canals that converge on KAH0204/32; NIWA18217, NIWA18218, NIWA 18219 the oscule (Fig. 5A). A fringe of very long diactines is and NIWA18220 from NIWA Stn KAH0204/40. present on the base of some specimens. NIWA Stations: E731 (NIWA25613); KAH0204/02 Dimensions 50 (20–65) mm high, 50 (20–80) mm (NIWA18225); KAH0204/07 (NIWA18226); wide, and 40 (20–60) mm thick. KAH0204/29 (NIWA18227, NIWA18228, NIWA18229, Texture stony overall, but crumbly on the surface, NIWA18230); KAH0204/32 (NIWA18231, NIWA18232, ectosomal region easily sloughed off. NIWA18233); KAH0204/38 (NIWA18234); Surface smooth, slightly rough, unabraided sections KAH0204/40 (NIWA18235, NIWA18236); KAH0204/44 have short robust conules dispersed about 5–10 mm (NIWA18237); KAH0204/47 (NIWA18238); apart, raised from projecting diactines (Fig. 5C). RAPUHIA2-15 (25647); T226 (NIWA25650); Colour in ethanol pale orange (5A3), honey yellow TAN0104/001 (NIWA18240); TAN0104/002 (5D6) to topaz (5C5). (NIWA1824, NIWA18247, NIWA18248); TAN0104/048 Choanosomal skeleton composed of robust smooth (NIWA18239); TAN0104/148 (NIWA18242); tetraclones tightly zygosed with adjacent desmas both TAN0104/194 (NIWA18243); TAN0104/288 terminally and laterally. Long diactines emerge from (NIWA18244); TAN0104/289 (NIWA18245); the centre of the sponge in tracts, piercing the surface TAN0104/394 (NIWA18246); TAN0107/232; X201 in low conules. Large microscleres are abundant and (NIWA18223); Z9025 (NIWA18224). regularly distributed within the choanosome. Ectosomal skeleton an encrustation of small micro- Type locality: Bay of Plenty, North Island, New Zea- scleres. land. Megascleres (Table 4) tetraclone desmas with very thick smooth clones that rarely branch, and if they do Distribution: Seamount east of Three Kings Ridge; it is towards the terminus of the clone (Fig. 5D). The West Cavalli, Cavalli, South Cavalli Seamounts, and clones are ornamented with single or double pairs of Seamount 441, northeastern New Zealand; Knights tubercles or spires of numerous tubercles. Spires oc- Terrace, East of Poor Knights Islands; Rumble V cur around the axis of the clone (which is generally Seamount, West of White Island, and Ngatoro Ridge, quite smooth) and are generally in the most mature Bay of Plenty; North of Raoul Island, Kermadec Ridge; spicules (Fig. 5E). Zygosis is terminal although zygoses Diabolical, Graveyard, Morgue, and Scroll Seamounts, frequently adhere laterally with a saddle-shaped for- Chatham Rise. mation when against adjacent clones (Fig. 5D). The zygoses in older portions of the skeleton are often so Habitat: Attached to small pieces of scleractinean coral intricate and over-developed as to be enlarged many rubble and rock surfaces on the sides of seamounts. times thicker than the clones. Dichotriaenes with one Many of the specimens examined appear to have set- set of clads shorter than the other two, and one of these tled on hard coral, eventually engulfing them as they

Table 4. Spicule dimensions (µm) of Neoaulaxinia persicum n. sp. Desma (Clone L/T) Cladome (Max ø) Diactines Ms 1 (L) Ms 2.1 (L) Desma (Overall size) Rhabd (L) (L) Ms 2.2 (L)

New Zealand NIWA18215 250–500/60–100 627 (350–875) 750+ 17 (15–20) 50 (43–65) (holotype) 675–750 496 (375–550) 26 (18–35)

NIWA18216 375–450/100–120 694 (525–900) 800+ 19 (18–23) 46 (33–55) (paratype) 750–750 388 (375–400) 21 (18–32)

NIWA18221 250–500/80–100 600 (475–700) 1250+ 17 (15–20) 46 (37–50) (paratype) 500–875 — 24 (20–25)

21 shorter clads is usually shorter than the other giving • The immature sponge could be mistaken for Ne- the cladome a lopsided appearance (Fig. 5F). The re- osiponia motukawanui n. sp. maining two sets of clads curve inwards, sometimes acutely, to resemble calipers. The tips of the clads are Neosiphonia Sollas, 1888 often blunt and strongylote. The rhad is often curved with a strongylote end. Diactines (Fig. 5D) are very thick with oxeote or frequently strongylote ends and Club-shaped to spherical body with a restricted base are up to several mm long. They are bundled in tracts of attachment, a bundle of vertical canals arises in the that emerge within the choanosome. Mature spicules central axis of the sponge emerging within a cavity on are over 3 mm long. the sponge apex, the cavity is covered with a delicate Microscleres (ectosomal) are small strongyloam- parchment-like ectosomal roof; desmas are branched phiasters with very robust rays (Table 4, Ms 1), cho- and ramified tetraclones with sculpted sinuous or tu- anosomal microscleres are large streptasters (Fig. 5G) berculate projections on the clones. Zygosis is intricate with a reduced number of rays (Table 4, Ms 2.1), and in inner parts of the skeleton, weak in the outer (grow- smaller streptasters with numerous sharp rays (Table ing) regions of the sponge; dichotriaenes are generally 4, Ms 2.2). short-shafted (100–200 µm on average), the cladome is wide with clads of irregular length; microscleres are Etymology: persicum (L.) = peach. acanthose in two clearly differentiated groups, a robust strongyloamphiaster restricted to an ectosomal crust, Remarks: Neoaulaxinia persicum is clearly differentiated and a choanosomal streptaster with conical rays. morphologically from N. clavata and N. zingiberadix. The new species is egg-shaped with a restricted base, Type species: Neosiphonia superstes Sollas, 1888. contrasting sharply with the elongate club-shaped N. clavata and tuberose N. zingiberadix. The tracts of ox- Neosiphonia superstes Sollas, 1888 (Fig. 6) eas that emerge in a conule at regular intervals on the surface of N. persicum are also highly characteristic of Neosiphonia superstes Sollas, 1888: 299, pl. 31, figs 7–12; Lévi sponges that have been recovered with little damage. & Lévi 1983: 119, fig. 10; Lévi & Lévi 1988: 47, pl. 3, fig. At the microscopic level there are several clear 3; Pisera & Lévi 2002k: 366, figs 6–9. differences in the size of the various categories of mi- Material examined: croscleres and the morphology of the key diagnostic megascleres. In particular, the ornamentation of the NIWA Stations: E731 (NIWA25614); KAH0204/07 tetraclone desmas with multituberculate spires is (NIWA18258); KAH0204/32 (NIWA18257); unique in the genus and the clones are much thicker KAH0204/47 (NIWA18256); TAN0107/124 than in both other species. Zygosis is highly developed (NIWA18255); TAN0205/20 (NIWA25643); in mature spicules, with an enlarged complex tangle TAN0205/32 (NIWA18249, NIWA18250, NIWA18251); of terminal clasps and lateral saddle formations. The TAN0413/120 (NIWA25051). dichotriaenes in N. persicum have a much wider and more irregular cladome and a longer rhabdome than Distribution: Challenger Station 173, off Matuka, Fiji in the other two species. The choanosomal microscleres Islands (holotype); BIOCAL Stn DW66, south New are the largest and most robust of species in the genus Caledonian slope of Norfolk Ridge; East Kermadec and all microscleres are only very slightly acanthose, Ridge Slope, Volcano E, Kermadec Ridge; Rumble III much less so than all other Phymatellidae described Seamount and Mahina Knoll, Bay of Plenty; Seamount thus far. 441, West Cavalli, and Cavalli Seamounts, northeastern N. persicum is an easily recognised species that is so New Zealand. far known only from the north and east of New Zealand from the Three Kings Rise and the Chatham Rise. Habitat: Attached to hard dead coral or rocky substratum. Depth range 430–999 m.

Key diagnostic characters: Description: • Sponge is spherical to egg-shaped with an apical Morphology short, squat, spherical body elevated on a oscule and subdermal canals short restricted base of attachment with a moderately • Desmas with very thick smooth clones, with spires deep apical cavity (Fig. 6A). of tubercles, zygosis is very intricate and robust Dimensions typically 35 (20–50) mm high and 35 • Microscleres are very large and only slightly acan- (15–60) mm at the widest part of the body. The base those ranges from 15–20 mm wide. The apical cavity indents 30–35% of the apex and ranges in depth from 5–10 mm.

22 Figure 6. Neosiphona superstes Sollas, 1888.

23 In macerated specimens the grouped vertical canals phiasters. The microscleres were originally described that run through the centre of the sponge are visible, as ‘spirasters’ by Sollas (1888) and Pisera and Lévi but in complete specimens the canals are covered with (2002k), but are clearly illustrated as amphiasters in the a loose flaky ectosome. former publication, and are illustrated in two categories Texture firm, almost stony. (strongyloamphiasters and amphiasters) in Pisera and Surface irregularly mounded to smooth. Lévi (2002k) as in other members of the Phymatellidae. Colour in life ivory (4B3), in ethanol honey yellow It is likely that Sollas did not find the robust strongy- (5D6). loamphiasters in the holotype as they are restricted to Choanosomal skeleton composed of highly branched an ectosomal crust, which would have been absent in tetraclones only weakly articulated with adjacent spi- the “deciduous” specimen. cules. Microscleres are abundant. Oxeas arise in tracts Apart from N. superstes, only two other species are from within the deeper choanosome but do not pierce known. N. fruticosa (Wilson, 1925) is a semi-planar the surface. branching mass 55 mm high, 90 mm wide, and 50 mm Ectosomal skeleton an encrustation of strongyloam- thick, of cylindrical, slightly flattened branches about phiasters. 7 mm diameter. It was first described from the Phil- Megascleres (Table 5) tetraclone desmas with smooth ippines. N. schmidti Sollas, 1888 from Havana, Cuba, uneven clones that bifurcate several times, each clone probably belongs to a different genus according to reducing rapidly in thickness, with tubercular and Pisera and Lévi (2002k). sculpted sinuous protuberances, especially abundant at the clone terminus (Fig. 6B). Zygosis can be weak Key diagnostic characters: in immature spicules but is generally relatively strong • Sponge is a short squat spherical body elevated on a and situated laterally and terminally. Dichotriaenes short stem with a broad apical depression in which have a short rhabd (Fig. 6C) and comparatively wide many canal openings are visible. cladome. The protoclad leading to the two deuteroclads • Desmas are highly branched with sinuous sculpted is comparatively long and the clads are separated by a zygoses. wide angle. Diactinal spicules are fusiform with long • Short rhabds on the dichotriaenes (well under 300 fine tapering ends. µm). Microscleres (Table 5) (ectosomal) are small • Microscleres are two types only (but these are in strongyloamphiasters with short robust rays (Table two sizes). 5, Ms 1, Fig. 6D, choanosomal microscleres are larger • The immature sponge could be mistaken for Neo- amphiaster/streptasters (Table 5, Ms 2, Fig. 6D, with aulaxinia clavata. fine oxeote rays. Neosiphonia motukawanui n. sp. (Fig. 7) Remarks: The distinguishing characteristics of this well- described species (Sollas 1888; Lévi & Lévi 1983, 1988; Material examined: Pisera & Lévi 2002k) is the rounded stalked body with Type material: Holotype NIWA18261 and paratype an apical cavity and siphon-like central canals, and at NIWA18259 from NIWA Stn KAH0204/38. the microscopic level the morphology of the desmas NIWA Stations: KAH0204/29 (NIWA18265, (strongly branched with sculpted sinuous tuberculate NIWA18266, NIWA18267); KAH0204/32 (NIWA18264); projections) and the large size of the streptaster/am- KAH0204/38 (NIWA18268, NIWA18269, NIWA18270,

Table 5. Spicule dimensions (µm) of Neosiphona superstes Sollas, 1888. Desma (Clone L/T) Cladome (Max ø) Diactines Ms 1 (L) Ms 2 (L) Desma (Overall size) Rhabd (L) (L)

Fiji BMNH 1891.5.4.11 100–200/70 550 2,280 — 45 (holotype) 150 New Caledonia Lévi & Lévi (1983) 200–350/70 400 900 (min) — — 600–700 100 New Zealand NIWA18250 100–375/60–80 348 (250–500) 750 (min) 16 (13–18) 32 (25–43) 500–600 136 (75–200) NIWA18249 150–400/50–80 433 (375–525) 675 (min) 19 (18–20) 40 (35–45) 500–700 200 (150–275)

24 Figure 7. Neosiphona motukawanui n. sp.

25 NIWA18271, NIWA18272); KAH0204/47 (NIWA18260); Ectosomal skeleton a crust of robust strongyloam- KAH0204/52 (NIWA18262, NIWA18263, NIWA18273, phiasters. Oxeas arise in tracts from within the deeper NIWA18274, NIWA18275). choanosome, piercing the surface, and in some loca- Type locality: West Cavalli Seamount, Northland. tions forming a distinct beard. Megascleres (Table 6) tetraclone desmas with very Distribution: Cavalli Seamount Region, northeastern thick short clones (Fig. 7C), tuberculation variable and New Zealand. more prominent on the ends of the clones. Single, multiple and spires of tubercles are common. Clones divide Habitat: Sponges were dredged from hard rocky sub- at least once, and often twice. Zygoses are very strong, strata or were attached to small pieces of coral. Depth intricate, and enlarged. Dichotriaenes (Fig. 7D) with range 780–910 m. very wide cladome; clads are thick, irregular and often of uneven length, rhabd short and often strongylote. Description: Dichotriaenes of NIWA18261 are frequently trichotri- Morphology a block-shaped mass with a restricted base aenose and the clads are occasionally strongylote, as of attachment, the apex of which is a flattened parch- is the rhabd. Diactines are very long, straight, evenly ment-textured roof, under which numerous aquiferous thick and strongylote. canals open from within the sponge (Fig. 7A,B). The Microscleres (ectosomal) are robust strongyloam- upper third of some specimens may be expanded to phiasters with a few blunt conical rays (Table 6 Ms form a shallow cup with thick rolled edges (Fig. 7B). 1; Fig. 7E), choanosomal microscleres are larger The base of these sponges is very narrow and root-like amphiaster/streptasters with fine rays (Table 6 Ms (Fig. 7A, B). NIWA18268 has a thin beard of diatines 2; Fig. 7E). In addition to the standard microscleres, about 40 mm long extending from around the base of specimen NIWA18264 has rhabds with short projec- the sponge into the substratum, probably acting as an tions postioned very irregularly along the length of the anchoring device. Immature sponges are elongate to spicule which is 17–20 µm long. These were not seen spherical with a shallow broad parchment-like apex, in other specimens, and are most probably malformed gradually thinning to a narrow base of attachment. The strongyloamphiasters. sponges are generally taller than they are wide. Dimensions of immature specimens in the collection Etymology: Named for the largest island in the Cavalli are 25 mm high, 15 mm at the widest point, and 5 mm Island group, Motukawanui, encompassed by the Cav- at the base. Most specimens are approximately 50 mm alli Seamount region that is the general type locality high, 35 mm wide (maximum), and 15 mm wide at the of this new species. In the literal sense, motu means base. Mature specimens are 60–80 mm high, 40–60 mm ‘island’, kawa is the New Zealand pepper tree used at the widest point, and have a base of 20 mm. extensively in ceremonial protocol, and nui means ‘big’ Texture internally incompressible, externally flakey, (motukawanui, New Zealand Māori). crumbly, easily abraided, especially at the apex. Surface typically smooth, flakey or crumbly. The Remarks: Neosiphonia motukawanui is only the fourth surface of many specimens is tufted with long diactinal known species of this genus, and one of these, spicules that form conules on the surface, but these are N. schmidti Sollas, 1888 from Havana, Cuba, is thought often abraided away in damaged specimens. to belong to a different genus (Pisera & Lévi 2002k). Colour apricot yellow (5B6) in ethanol. Neosiphonia is characterised primarily by a massive Choanosomal skeleton dominated by desmas with globular body that narrows to a restricted columnar thick tuberculate clones and large intricate zygoses. base of attachment, the apex of which consists of a cov- Oxeote microscleres are abundant. ered shallow depression or atrium into which vertical aquiferous canals open. Table 6. Spicule dimensions (µm) of Neosiphonia motukawanui n. sp. Desma (Clone L/T) Cladome (Max ø) Diactines Ms 1 (L) Ms 2 (L) Desma (Overall size) Rhabd (L) (L)

New Zealand NIWA18261 125–375/70–80 523 (450–675) all broken 18 (18–20) 30 (23–35) (holotype) 460–600 138 (75–250)

NIWA18259 250–400/90–100 619 (425–900) 600+ 21 (18–28) 33 (20–43) (paratype) 500–800 142 (75–250)

NIWA18264 250–500/70–100 590 (350–725) all broken 18 (18–20) 31 (25–38) 500–750 100

26 Neosiphonia motukawanui differs from the genus type Material examined: N. superstes Sollas in the shape of the sponge; the body NIWA Stations: B314 (NIWA18277); E731 (NIWA25612); region of N. superstes is more globular with a columnar KAH0011/30 (NIWA18278, NIWA18279); KAH0011/41 ‘stalk’ (see Fig. 6A), whereas the body of the former (NIWA18280); KAH9907/49 (NIWA18281); species tapers to a thin attenuating stalk (see Fig. 7A). TAN0413/099 (NIWA25056); TAN0413/112 The apical depression atop N. motukawanui is extremely (NIWA25055); TAN0413/117 (NIWA25058); prominent, covering the entire flattened apex of the TAN0413/118 (NIWA25057); X861 (NIWA25622). sponge, while in N. superstes the atrium is smaller and IRD Stations: LITHIST DW7 (NIWA18486, deeper. Long tufted diactines in N. motukawanui are NIWA18487). also a prominent feature in well-preserved material. The dichotriaenes have a wider, larger cladome than Distribution: BIOCAL Stn DW 66 (holotype) and in N. superstes and the strongyloamphiasters are much Éponge Seamount, south New Caledonian slope of thicker and slightly bigger. The desmas of N. motuka- Norfolk Ridge; Eastern edge of the Challenger Plateau; wanui are more robust and tuberculate than the thinner, west of White Island and Rungapapa Knoll, Bay of less ornamented desmas of N. superstes. Plenty; South Kermadec Ridge. Younger specimens of N. motukawanui may be mistaken in the field forNeoaulaxinia persicum, but the Habitat: Growing on hard volcanic rock; the cover latter are spherical without an atrium over the apical photo of this memoir shows the surface of a boulder cavity, and the former are always club-shaped with from a volcanic seamount, clearly indicating the earliest low blunt conules. The nature of the ornamentation stages of settlement and growth of this species. Settle- of the desmas, the size and shape of the microscleres, ment and coalescence of adjacent individuals indicates and the greater rhabd length of the dichotriaenes in N. that older sponge masses may be made up of several persicum, are always indicative, however. individuals. Depth range 141–1132 m.

Key diagnostic characters: • Sponge is block-, club-, or shallow cup-shaped and Description: the flattened top of the sponge is a delicate ‘parch- Morphology of mature sponges a folded, tubular mass ment’ over a shallow cavity; the bottom of the cavity (Fig. 8A). The smallest individuals are rounded tu- has numerous aquiferous canals opening into it. berculate nodules with a slightly restricted base and a • Desmas are very thick, branched several times, and deep cavity on the apex. As these sponges mature, the are tuberculate, with single or double spires. walls extend upwards and outwards, the sponge then • The robust microscleres are very thick with few resembling a low flat, or high narrow chalice (Fig. 8B; conical blunt rays. see also cover photo). Portions of large sponges can • Could be confused with Neoaulaxinia persicum. appear to be lamellar or tubular when broken. Dimensions of juveniles 5 mm high, 4 mm wide at Reidispongia Lévi & Lévi, 1988 apex, 4 mm diameter at base; typical immature chalice- shaped sponges 15–22 mm high, 20 mm wide at apex, Cup-shaped with thin smooth walls, elongate chalice- 9 mm wide at base; the holotype is 60–80 mm high, shaped as a young sponge, foliose as a mature sponge; 90–100 mm wide at the apex, but the sponge grows tetraclone desmas are highly ramified with frequently much larger, up to 150 mm high and 300 mm wide; smooth branched clones and tuberculate termini. Zy- 3–5 mm thick. gosis is weak or non-existant on the outer (younger) Texture stony to touch, but the lamella is easily portions of the lamellae, otherwise it is intricate and broken and crushed. very strong in the older skeleton; microscleres are Surface very smooth, slightly glistening owing to two types of amphiaster in two closely related size faintly projecting diactines, oscules not visible. categories, one a strongyloamphiaster, and the other Colour characteristically vivid blue (22B8) in life, an amphiaster with fine rays. this colour able to be retained in ethanol, becoming sapphire blue (23D7), but several sponges are cream Type species: Reidispongia coerulea Lévi & Lévi, 1988. (4A3) or putty-coloured (4B2) in preservative. Choanosomal skeleton very regular and packed with Reidispongia coerulea Lévi & Lévi, 1988 (Fig. 8) small tetraclone desmas between which are abundant amphiasters. Dichotriaenes are regularly spaced along Reidispongia coerulea Lévi & Lévi, 1988: 245; Pisera & Lévi the ectosome with rhabds clearly visible perpendicular 2002k: 371, figs 11–14; Schlacher-Hoenlinger et al. 2005: to the ectosome surface. Slightly sinuous diactines are 688, figs 4D, 20, 34. dispersed sparsely throughout the outer choanosome (Fig. 8C). These frequently pierce the ectosome.

27 Figure 8. Reidispongia coerulea Lévi & Lévi, 1988.

28 Ectosomal skeleton a crust of strongyloamphiasters. Remarks: This sponge has been well described by Lévi Megascleres (Table 7) desmas (Fig. 8C,D) are smooth, and Lévi (1988) and Pisera and Lévi (2002k). The New highly branched and ramified, branching up to 4 times, Zealand specimens are almost identifical in all aspects and the termini are tuberculate. The centre of the lamel- of morphology and spicule dimensions. The descrip- la zygosis is very intricate and forms a rigid enlarged tion given here adds to knowledge of the settlement articulation mass (Fig. 8D); in the outer (younger) patterns and morphology of the most juvenile forms. sections of the lamella, zygosis is weaker (Fig. 8C) or non-existant. Dichotriaenes have long straight deute- Key diagnostic characters: roclads and a relatively short spured rhabd (Fig. 8E). • Folded thin-walled vivid blue mass or tiny blue Diactines (Fig. 8C) are thickened centrally, thinner and chalices or nubbins. flexuous on the ends that are strongylote. • Two clearly differentiated types of amphiaster Microscleres (ectosomal) are robust strongyloam- microscleres of a similar size. phiasters (Table 7 Ms 1); choanosomal microscleres are larger fine-rayed amphiasters (Table 7, Ms 2; Fig. 8F).

Table 7. Spicule dimensions (µm) of Reidispongia coerulea Lévi & Lévi, 1988. Desma (Clone L/T) Cladome (Max ø) Diactines Ms 1 (L) Ms 2 (L) Desma (Overall size) Rhabd (L) (L)

New Zealand NIWA 18277 200–300/20–30 300 (220–400) 446 (250–770)+ 8 (8–11) 15 (13–18) 300–500 200 (160–270)

New Caledonia Lévi & Lévi (1988) 200/20 300 500–1000 9–10 15–18 300–475 125–200/18–20

S.-H. et al. (2005) —/20–30 190–300 500–1000 10–13 16–21 380-420 —

Family CORALLISTIDAE Sollas, 1888 the possession of two types of spirasters (with short blunt and long oxeote rays), a very large streptaster in Corallistidae Sollas, 1888: 301 (partim); Topsent 1894: 290 some species, and smooth anisoxeas; Corallistes also has (partim); Schrammen 1910: 64 (partim). two types of spiraster, both with oxeote rays (short and long rays), but lacks the anisoxeas. Research here has Massive ear-, vase-, plate-, or cup-shaped lithistid Dem- resulted in the transfer of C. fulvodesmus Lévi & Lévi, ospongiae; choanosomal desmas are dicranoclones, 1983 to Neoschrammeniella as it possesses anisoxeas ectosomal megascleres are dichotriaenes, microscleres and a very large streptaster, amongst other characters. are spirasters typically of two types; streptasters, am- C. multituberculatus Lévi & Lévi, 1983 is probably also phiasters, microxeas, microstyles, and microstrongyles a species of Neoschrammeniella as it has two types of (from Pisera & Lévi 2002c). spiraster, while C. undulatus Lévi & Lévi, 1983 has only one and remains in Corallistes. Only two species Remarks: Redefinition of the Corallistidae by Pisera and of Corallistes are presently recognised in the Southwest Lévi (2002c) and description of new species of Coral- Pacific —C. undulatus Lévi & Lévi, 1983 and C. australis listes, Herengeria, and Neoschrammeniella by Schlacher- Schlacher-Hoenlinger, Pisera & Hooper, 2005. Hoenlinger et al. (2005) have facilitated the recognition Species of Herengeria are separated from Neoschram- of corallistid genera in New Zealand waters. Species meniella in the possession of acanthose centrotylote of Corallistidae occur in deep-water worldwide, but oxeas and spiral or rod shaped micropolyrhabd mi- Herengeria and Neoschrammeniella are so far known croscleres in addition to normal spirasters, and in the only from the Southwest Pacific. Most species are manner in which the desmas zygose. Awhiowhio n. poorly known and badly in need of revision (Pisera gen. is similar to Herengeria, but lacks the ectosomal & Lévi 2002c). acanthose microxeas and choanosomal spirasters, and Neoschrammeniella Pisera & Lévi (formerly Iouea the desmas are less ornamented with smooth nodular Lévi & Lévi, 1988) differs from Corallistes Schmidt in projections rather than multituberculate fungiform

29 projections. This new genus is endemic to New Zea- IRD Stations: LITHIST DW7 (NIWA18488). land waters. The separation of species with nodulose dichotri- Distribution: Éponge Seamount, and south New aene cladomes from Corallistes to Neophrissospongia, Caledonian slope of Norfolk Ridge (holotype); South suggested some years ago by Lévi (1991) and supported Norfolk Basin; Norfolk Ridge south of Norfolk Island; by Kelly (2000), further differentiates the family. This north, west and east of Three Kings Ridge, northern genus is represented in the southwest Pacific by a New Zealand; West and South Cavalli Seamounts, single species, Neophrissospongia microstylifer (Lévi & northeastern New Zealand; Rungapapa, Tuatoru and Lévi, 1983) (Pisera & Lévi 2002c), which has not been Mahina Knolls, Bay of Plenty, New Zealand. found in New Zealand waters. It is interesting to note that microfossil ectos- Habitat: Sponges were dredged from hard rocky sub- omal triaenes indistinguishable from those of extant strata; specimens frequently had coral incorporated Corallistidae were found in the Tutuiri Greensand into their bases. Depth range 180–1680 m. (Teurian–basal Waipawan) in the Chatham Islands (Buckeridge & Kelly 2005), and illustrated from the Description: Oamaru Diatomite at Oamaru in North Otago (Hinde Morphology a shallow cup-shape with flared margin & Holmes 1892, pl. 13, figs 7–8, 11–12), suggesting a and narrow base of attachment, or an erect concave more southern distribution for Corallistidae during plate attached along a restricted curved base (Fig. the Paleogene. 9A–C). Immature specimens form a nodule on the substratum with a single apical channel that gradu- Neoschrammeniella Pisera & Lévi, 2002 ally widens and expands as the sponge matures into either a cup or a plate. Several small specimens had Iouea Lévi & Lévi, 1988. hermitcrabs living in their narrow aperture. Dimensions of cups are 40–90 × 30–90 mm wide and Shallow cup-shaped or deep vase-shaped Corallistidae 25–50 mm high with a margin 4–8 mm thick. The base with a smooth surface; desmas are heavily tuberculate of attachement is circular or elliptical and varies with dicranoclones. Ectosomal spicules are smooth dichotri- the size of the sponge. Small sponges typically have a aenes. Medium-length anisoxeas with attenuated tips base of c. 10 mm; larger sponges have a base c. 40 mm may be present in the ectosome in several species and wide. Plate sponges are 25–120 mm wide and 25–60 large diactines may form a fringe along the sponge mar- mm high with a base of attachment 10–40 mm wide. gin. Microscleres are two types of acanthose spirasters, Texture very rigid, stony. one with robust rays and one with thin pointed rays, Surface smooth, granular to the touch, regular. and a large amphiaster/streptaster with smooth thin Colour cream (4A3) or putty-coloured (4B2) in etha- pointed rays in several species (emended from Pisera nol preservative. & Lévi 2002). Choanosomal skeleton dense, with very heavily tu- berculated dicranoclones (Fig. 9D). Type species: Iouea moreti Lévi & Lévi, 1988. Ectosomal skeleton a dense layer of robust microscleres that obscure the cladomes of the dichotriaenes. Neoschrammeniella fulvodesmus (Lévi & Lévi, Large streptasters are abundant in the choanosome with smaller thin-rayed spirasters (Fig. 9D). 1983) (Fig. 9) Megascleres (Table 8) — desmas are highly tu- Corallistes fulvodesmus Lévi & Lévi, 1983: 110, pl. 5, fig. 2,5, berculate dicranoclones with fungiform tubercles pl. 13, 2, fig. 4. across the primary arched clone (Fig. 9E–F). There are approximately 4 clones arising off each arch. Di- chotriaenes are robust with short conical clads (Fig. Material examined: 9F), and characterised by bifurcating tips, a character NIWA Stations: I92 (NIWA18289); KAH0204/29 first noted by Lévi and Lévi (1983). The diactines are (NIWA18295); KAH0204/32 (NIWA18293); extremely long. Attenuated oxeas are present in ecto- KAH0204/40 (NIWA18292); KAH0204/44 somal preparations. (NIWA18294); S571 (NIWA18291); S572 (NIWA18284, It is worth noting that in specimen NIWA18298 NIWA18285); TAN0413/070 (NIWA25049); from Rungapapa Knoll, Bay of Plenty, over 50% of the TAN0413/111 (NIWA18296, NIWA18297, dichotraienes are severely malformed, with only two NIWA25050); TAN0413/173 (NIWA18298); U606 of the three clads present, with a rhabd-like structure (NIWA18287); X128 (NIWA18283); X140 (NIWA18290); continuing through the cladome, to odd spurs emerg- Z9025 (NIWA18288); Z2098 (NIWA18286); Z9751 ing from the centre of rhabds. The tips of cladomes (NIWA18515).

30 Figure 9. Neoschrammeniella fulvodesmus (Lévi & Lévi, 1983).

31 Table 8. Spicule dimensions (µm) of Neoschrammeniella fulvodesmus (Lévi & Lévi, 1983).

New Zealand New Caledonia NIWA18285 NIWA18288 Lévi & Lévi (1983)

Desma Clone L/T 200/50–120 100–200/60–100 —/100–150 Overall size 400–600 500–-600 600 Cladome (Max ø) 500 (400–625) 348 (250–400) 80–200 Rhabd (L) 500 (400–625) 475 (375–625) 600 Diactine (long) > 1 mm > 1 mm 1200–1300 Anisoxea — 396 (240–600) — Spiraster (robust) 17 (15–20) 19 (15–25) 10–15 Spiraster (thin) 21 (15–25) 22 (20–25) not measured but figured Streptasters 61 (52–67) 58 (50–62) 35–45 Ray length 28 25–30 20–25 are frequently bifurcate, but in this specimen they are N. moreti and N. fulvodesmus (both have very strongly often trifurcate and multiply branched. tuberculate desmas with terminal articulation), and Microscleres (Table 8) (ectosomal) are robust thus the decision only to expand the diagnosis is spirasters (Fig. 9H); choanosomal microscleres are upheld. spirasters with numerous thin rays, and very large streptasters with thin rays (Fig. 9G). Key diagnostic characters: • Relatively thin-walled regular cup or plate with Remarks: New Zealand specimens examined here are smooth granular surface. clearly conspecific with Corallistes fulvodesmus Lévi & • Diagnostic spiraster microscleres of two types Lévi, 1983, which is now included in Neoschrammeniella. (thick-rayed robust spirasters and very large thin- Corallistid genera are largely differentiated on the rayed spirasters). morphology of the microscleres and the presence of • May be mistaken in the field forHerengeria vasiformis various diactines in the ectosome. Neoschrammeniella but the concave surface is much less furry; the norfolkii Schlacher-Hoenlinger, Pisera & Hooper, 2005, microscleres of H. vasiformis are also unique, and N. antarctica n. sp. (described below), and New Zealand diagnostic of that species. and New Caledonian specimens of N. fulvodesmus • May be confused in the field withAwhiowhio osheai are characterised by the presence of extremely large n. sp., but this sponge has raised oscules on the streptaster/ amphiasters in the choanosome in addition concave face, unique polymicrorhabds, and lacks to two forms of spiraster, which, in combination with the large streptaster category. anisoxeas, define the genus. This cluster of characters notwithstanding, the large streptaster/amphiasters Neoschrammeniella antarctica n. sp. (Fig. 10) are present in all species except the actual type of the genus, i.e. N. moreti (Lévi & Lévi, 1988), and N. castrum Material examined: Schlacher-Hoenlinger, Pisera & H, 2005. Two courses Type material: Holotype NIWA18299 and paratype of action are available to accommodate these species NIWA18300 from NIWA Stn A0527. with an additional huge streptaster category. Either a new genus can be erected for these sponges with an Type locality: Ross Sea, Antarctica. additional microsclere category, or the diagnosis of Neoschrammeniella can be emended. Although there Distribution: Ross Sea, Antarctica. may seem to be some justification for the former action (N. moreti is columnar with an axial canal compared to Habitat: Both specimens were dredged from hard the cup- or plate-shaped morphology of N. fulvodesmus), substrata; the smaller of the two bears remains of the N. antarctica forms a tall narrow cup that approaches substratum, which appears to be igneous rock. Depth the morphology of the genus type. Moreover, there range 352 m. is very little difference in the desma morphologies of

32 Figure 10. Neoschrammeniella antarctica n. sp.

33 Description: Table 9. Spicule dimensions (µm) of Neoschrammeniella Morphology of holotype is a tall, thick-walled, regular antarctica n. sp. conical cup with a deep aperture and square margin New Zealand at the apex (Fig. 10A). The cup diameter reduces to a NIWA 18299 (holotype) regular narrow base of attachment that is circular in profile. The smaller immature paratype has a similar Desma Clone L/T 350/70 morphology but the sponge has rounder sides and Overall size 350–750 the upper margin is rounded. There is no aperture, just a shallow depression with a soft parchment-like Cladome (Max ø) 386 (250–500) surface. Rhabd (L) 612 (425–725) Dimensions of holotype: 50 × 60 mm wide at apex, > 2 mm tapering to 20 mm wide at base, 65 mm high, wall Diactine (long) thickness 16 mm, aperture 30 long × 15 mm wide. Anisoxea 203 (150–350) Paratype 34 × 30 mm wide at apex, tapering to 12 mm Spiraster (robust) 14 (8–18) wide at base, 33 mm high, central cavity 31 × 13 mm wide, 2 mm deep. Spiraster (thin) 17 (15–20) Texture stony. Streptasters 74 (57–90) Surface smooth, slightly pitted, granular, except Ray length 25–38 for the concave surface of the aperture that is furry with projecting diactines, and the inner margin that is fringed in places with the same spicules. are much larger overall. The desmas of N. fulvodesmus Choanosomal skeleton extremely dense with large have very large nodular fungiform tubercles that are heavily tuberculate dicranoclones and abundant mi- commoner on the primary arched clone, as opposed croscleres. The choanosome is traversed by extremely to being spread regularly over all clones in N. norfolkii. long thick diactines. The desmas of N. antarctica have large pedestals of Ectosomal skeleton crowded with robust spirasters tubercles as well, principally on the arched side of the and dichotriaenes. primary clone. Megascleres (Table 9) — desmas are large dicrano- While the dichotriaenes in all three species are quite clones with clones that are heavily ornamented on all similar in cladome shape, the rhab is longer than in the surfaces with bumpy fungiform tubercles (Fig. 10 B, two other species and the clads are frequently trifur- C). Zygosis is predominantly terminal, with clasp-like cate. All categories of microscleres are quite similar in structures. Dichotriaenes are large and robust with length with the exception of the thin-rayed spirasters short conical clads, the tips of which are frequently in N. norfolkii (22–26 um), which are larger than in the bifurcate (Fig. 10 B, C). The diactines are extremely other two species. The large category of streptaster long and thick. Oxeas with attenuated tips are present is quite a lot smaller in N. fulvodesmus, however. The in ectosomal preparations. anisoxeas of N. antarctica are smaller than those in N. Microscleres (Table 9) (ectosomal) are robust fulvodesmus; these spicules are not mentioned in the spirasters that form a crust (Fig. 10E); choanosomal description of N. norfolkii. microscleres are small spirasters with numerous thin N. antarctica is the first record of a lithistid sponge rays and large streptasters with thin smooth rays (Fig. from a polar region and the only lithistid sponge record 10D, E). south of the Challenger Plateau and the Chatham Rise of New Zealand. Etymology: Named for its Antarctic distribution. Key diagnostic characters: Remarks: Neoschrammeniella antarctica is very similar • Very thick-walled conical cup. to N. fulvodesmus from northern New Zealand and • From Antarctica. the Norfolk Ridge and to N. norfolkii from the Norfolk Ridge, but differs in gross morphology, in the subtle Herengeria Lévi & Lévi, 1988 detail of the morphology and ornamentation of the desmas, and in the size of the accessory megascleres. Massive or cup-shaped. Massive species have a lateral Specimens of N. antarctica are tall thick-walled vases inhalant sieve-pore area and long tufts of diactines; with a deep cavity and square lip, whereas both N. cup-shaped species have no localised inhalant struc- fulvodesmus and N. norfolkii appear more cup-shaped. ture, but the upper concave part of the cup is bristly. The desmas in all three species are quite similar in Choanosomal megascleres are smooth dicranoclones overall size but those of N. fulvodesmus and antarctica with multituberculate fungiform nodules and spires;

34 zygoses are intricate, terminal or predominantly lateral ‘auricle’ that is softer and more porous than the sur- via saddle-shaped formations, clones forming rein- rounding sponge material, representing a localised forced ‘bridges’ around ‘trabeculae’ or channels in the concentration of inhalant ostia. Tufts of very long network. Ectosomal spicules are smooth dichotriaenes; spicules emerge from the apex and sides of the sponge choanosomal microscleres are abundant acanthose (Fig. 11A, B). A short oscular chimney may emerge centrotylote microxeas and long-rayed streptaster/am- from base of sponge. phiasters. Ectosomal microscleres include short-rayed Dimensions 15–25 mm high, 8–20 mm wide, 6–15 spirasters, amphiasters, and highly irregular micropo- mm thick, excluding spicule tufts that emerge from lyrhabds. Large oxeas protrude from the upper surface 2–7 mm beyond the sponge surface. The holotype (modified from Pisera & Lévi 2002c). was described as 15 mm high and 10 mm wide (Lévi & Lévi 1988). Type species: Herengeria auriculata Lévi & Lévi, 1988. Texture stony; inhalant auricle is soft, crumbly. Surface relatively smooth except for the tufts of Remarks: The diagnosis has been modified from that spicules. given by Pisera and Lévi (2002c) to include a new form Colour apricot-yellow (5B6) in life and in ethanol, of choanosomal amphiaster microsclere found in all with a reddish tinge. specimens of Herengeria auriculata. Choanosomal skeleton composed of very large dicranoclone desmas that form a cavernous structure Herengeria auriculata Lévi & Lévi, 1988 (Fig. 11C), in between which are dispersed numerous microxeas, sharp-rayed spirasters and, less commonly, (Fig. 11A–G) long-rayed amphiasters. Enormous tracts of oxeas are Herengeria auriculata Lévi & Lévi, 1988: 250, pl. 5, fig. 5; visible emerging from the choanosome, penetrating Schlacher-Hoenlinger, Pisera & Hooper 2005: 665, figs and emerging beyond the sponge surface. 2G, 11, 25. Ectosomal skeleton a crust of microxeas and spiraled microscleres. Material examined: Megascleres (Table 10) dicranoclone desmas (Fig. 11 NIWA Stations: U591 (NIWA18301); Z2098 C,D), slightly curved, tripod-shaped, bearing abundant (NIWA18302). fungiform projections, the upper part of each being IRD Stations: LITHIST DW7 (NIWA18516). nodulose (Fig. 11D). Zygoses are terminal and lateral along the clones between the fungiform processes. Di- Distribution: BIOCAL Stn DW 66 (holotype), Éponge chotriaenes of typical form with a regular long conical and Introuvable Seamounts, south New Caledonian rhabd and regular smooth cladome. Diactines, very slope of Norfolk Ridge; Tui Seamount and north of long, centrally thickened oxeas that project in bundles Three Kings Ridge, northern New Zealand. from the choanosome to form tufts on the surface. Microscleres (ectosomal) acanthose microxeas, either Habitat: Free-living with no indication of an attach- acutely bent in the middle or straight and centrotylote ment. Depth range 486–920 m. (Fig. 11E,G); ectosomal spirasters, thick curved spirals (Table 10 Ms 1; Fig. 11E); choanosomal spirasters with Description: numerous sharp slightly curved rays (Table 10 Ms 2; Morphology globular, egg-shaped, with a truncate Fig. 11F); choanosomal amphiasters with long curved apex. One side of the sponge is a flattened demarcated rays (Table 10 Ms 3; Fig. 11F).

Table 10. Spicule dimensions (µm) of Herengeria auriculata Lévi & Lévi, 1988. Clone (thickness) Cladome Diactines Microxeas Ms 1 (L) Ms 2 (L) Ms 3 (L) Overall size (Max ø) (L) (L) Ray (L) Ray (L) of arch Rhabd (L) New Caledonia Lévi & Lévi (1988) 50–75 160–680 < 10 mm 70–100 23–33 20–22 – (holotype) 450–800 300–475

S.-H. et al. (2005) 50–75 245–500 10 mm 70–100 32–40 19–25 – 450–800 300–475

New Zealand NIWA18301 80–100 454 (375–625) < 10 mm 85 (75–95) 30 (25–35) 23 (18–25) 23 (20–25) 375–1000 366 (270–430) 5–6 10–13

35 Figure 11. Herengeria auriculata Lévi & Lévi, 1988.

36 Remarks: The New Zealand specimens conform well Dimensions 86 (45–120) x 79 (45–115) mm wide, to the holotype description. All New Zealand and 42 (20–80) mm high, wall thickness 6–10 mm (outer new New Caledonian specimens contain an addi- margin) to 12–25 mm at the base. tional choanosomal microsclere — an amphiaster with Texture stony. long, curved rays — which has not been described Surface — internal surface furry with projecting previously. The genus diagnosis has been emended diactinal spicules (Fig. 12B), surface textured, slightly accordingly. rough; convex outer surface rough and slightly irregular. Key diagnostic characters: Colour in life ivory (4B3), in preservative teak brown • Small stony nubbin sponges with tufts of long spi- (6F5). cules emerging from the top Choanosomal skeleton dense with aquiferous canals • With an ear-shaped expansion that contains local- visible under the ectosome, but the desma skeleton is ised inhalant pores largely obscured by extremely abundant acanthose microxeas, particularly on the outer ostial face. Microxeas Herengeria vasiformis Schlacher-Hoenlinger, are absent from the oscular face. Desmas form thick Pisera & Hooper, 2005 (Fig. 12) heavy arches within the deep choanosome and a less dense reticulation in the ectosome. Herengeria vasiformis Schlacher-Hoenlinger, Pisera & Hooper, Ectosomal skeleton a dense crust of micropolyrhabd 2005: 668, figs 2F, 10, 24. microscleres (Fig. 12F). Megascleres (Table 11) — desmas very thick, with Material examined: 2–4 clones, heavily ornamented with fungiform nodu- NIWA Stations: E636 (NIWA18309); KAH0011/30 lose tubercles (Fig. 12E). Zygosis is lateral and terminal, (NIWA18312); I372 (NIWA25781); KAH0011/40 building a very dense reticulation in the mature inner (NIWA18313); KAH0011/41 (NIWA18314); sections of the sponge; desmas have a granular brown KAH0011/44 (NIWA18310); KAH9907/48 core, immature desmas are sparsely tuberculate with (NIWA18316, NIWA18317); KAH9907/50 (NIWA18319, relatively smooth thin clones; dichotriaenes (Fig. 12F), NIWA18320, NIWA18321, NIWA18322, NIWA18323); regular, long rhabd and short thick clads, rhabd often S572 (NIWA18308); TAN0205/32 (NIWA18315); with a spur; oxeote diactines, smooth, very thick cen- TAN0413/074 (NIWA18327); TAN0413/077 trally, with whip-like extremities. (NIWA25047); TAN0413/085 (NIWA18326); Microscleres — micropolyrhabds (Table 11 Ms 1; TAN0413/112 (NIWA18328, NIWA25046); Fig. 12G), acanthose, extremely irregular, ranging TAN0413/117 (NIWA18324); TAN0413/118 from rod-shaped to nodulose, to spiraster-shaped with (NIWA18325); U594 (NIWA18307); U606 (NIWA18303, thick blunt nodules. Most spicules are non-uniform, NIWA18304); X207 (NIWA18305); Z9748 [NIWA18817 with nodules projecting randomly along one or both (0CDN6853-J)]. sides; streptasters (Table 11 Ms 2), extremely fine and IRD Stations: LITHIST DW7 (NIWA18517). irregular, with very long thin occasional rays, rare; microxeas, slender, curved, or slightly angled around Distribution: New Caledonian slope of Norfolk Rige; central tylote expansion; acanthose with short sharp east Kermadec Ridge slope; South Norfolk Basin; spines that curve inwards towards middle of spicule east of Hikurangi Plateau; West Three Kings Ridge; (Fig. 12E). Doubtless Bay, North Cape; Tuatoru and Rungapapa Knolls, and west of White Island, Bay of Plenty, New Remarks: The New Zealand specimens agree well Zealand. with the holotype recently described from seamounts off southern New Caledonia on the Norfolk Ridge Habitat: Dredged from hard rocky (volcanic) substrata. (Schlacher-Hoenlinger et al. 2005), except that the Depth range 154–1100 m. streptaster microscleres (Table 11, Ms 2) in the New Zealand specimens are extremely rare, and the sponges Description: are larger with a thinner wall. The micropolyrhabds Morphology a shallow regular spherical cup, the sides are unique in Corallistidae, and unusual in the lithis- shorter than wide (Fig. 12B), occasionally vase-shaped tid sponges. They are comparable to similar micro- (Fig. 12A,D); immature specimens short and dome- scleres in several new species of Neoschrammeniella shaped with central cavity (Fig. 12C); margins of cup (Schlacher-Hoenlinger et al. 2005) but this genus lacks smooth, wavy; sections of cup margin may be indented; the centrotylote microxea — an abundant microsclere base flanged. within Herengeria. Herengeria vasiformis is very different

37 Figure 12. Herengeria vasiformis Schlacher-Hoenlinger, Pisera & Hooper, 2005.

38 Table 11. Spicule dimensions (µm) of Herengeria vasiformis Schlacher-Hoenlinger, Pisera & Hooper, 2005.

Clone (thickness) Cladome Diactines Microxeas Ms 1 Ms 2 Overall size (Max ø) (L) (L) (L) (L) of arch Rhabd (L) New Caledonia S.-H. et al. (2005) 40–60 452–606 (broken) 83–111 13–17 15–21 (holotype) 400–600 162–209

New Zealand NIWA18303 50–80 220 (150–300) 1–5 mm 116 (100–130) 18 (10–23) 18 375–750 530 (375–750) NIWA18305 60–90 225 (150–300) 2–5 mm 91 (80–103) 20 (13–25) 32.5 300–625 630 (450–750) NIWA18307 50–80 238 (180–300) 1–5 mm 114 (100–125) 21 (15–38) 26 (25–28) 300–500 614 (400–875) morphologically from the type species H. auriculata in geria. The discovery of a second species with similar that it is cup-shaped rather than nubbin-shaped, re- characteristics from the South Fiji Basin (unda), and quiring an expansion of the diagnosis given by Pisera a third species from the Chatham Rise (sepulchrum), and Lévi (2002c). confirmed that these species form a group closely related to Herengeria. Key diagnostic characters: The simple desma morphology is similar to that in • Irregular micropolyrhabds on the surface. species of Corallistes (Pisera & Lévi 2002c; Schlacher- • Thick-walled cup with a hairy concave surface; Hoenlinger et al. 2005), but desmas of Corallistes do not can be confused in the field withCostifer wilsoni or typically form spires. The lack of microxeas and large Pleroma menoui, but these are more ear-shaped. streptasters also suggests affinity with the genusCoral - • May be confused with Neoschrammeniella moreti listes, but this genus is reserved for species with two microscopically, but this species lacks acanthose types of sharp-rayed spirasters. Moreover, Corallistes microxeas. sensu strictu has been recorded only from the New Caledonian section of the Norfolk Ridge (C. australis Awhiowhio n. gen. Schlacher-Hoenlinger et al. 2005 and C. undulatus Lévi & Lévi 1983). Massive cup- or plate-shaped Corallistidae. Choano- Awhiowhio is defined primarily by the smooth somal megascleres are smooth nodulose dicranoclones granular surface (indicating the lack of long hair-like with spires of smooth nodules, forming very robust diactines), the lack of distinctive acanthose microxeas intricate zygoses with ‘trabeculae’ through the entire found in Herengeria, and the presence of diacranoclones mass. Ectosomal spicules are smooth dichotriaenes. that have smooth nodules rather than multitubercu- Choanosomal microscleres are small fine-rayed late fungiform tubercles. The desmas are typically streptaster/spirasters. Ectosomal microscleres are corallistid, however, with a tripod to polypod shape irregular acanthose micropolyrhabds or irregular and saddle-shaped lateral zygomes. Like Herengeria, nodulose spirasters. Awhiowhio has only two types of asterose microscleres — the ectosomal polyrhabd and the choanosomal Etymology: Named for the spiral shape of the micro- streptaster/spiraster with multiple rays. scleres in this genus, but also relating to the general possession of spiralled or spiraster microscleres in Awhiowhio osheai n. sp. (Fig. 13) Corallistidae (āwhiowhio, New Zealand Maori noun for a ‘spiral’, ‘winding’, ‘whirlwind’ or ‘whirlpool’, or Material examined: adjective for ‘circuitous’). Type material: Holotype NIWA18332 from NIWA Type species: Awhiowhio osheai n. sp., here designated. Stn KAH9907/50; paratypes NIWA18333 from NIWA Stn E636, NIWA18334 from NIWA Stn KAH0011/30, Remarks: Awhiowhio n. gen. is established a Bay of NIWA25043 from TAN0413/093, and NIWA25042 Plenty species (A. osheai) that was initially taken to be from TAN0413/117. a species of Herengeria, but it lacks the characteristic NIWA Stations: KAH9907/48 (NIWA18335); acanthose ectosomal microxeas, nodulose tuberculate TAN0413/085 (NIWA25044); TAN0413/093 dicranoclones, and long hair-like diactines of Heren- (NIWA18336, NIWA25041); Z2364 (NIWA25619).

39 Figure 13. Awhiowhio osheai n. sp.

40 Type locality: Bay of Plenty, New Zealand. New Zealand deep-sea faunas and his passion for the discovery of new lithistid sponges and other inverte- Distribution: West of White Island, Rungapapa and brate groups. Tuatoru Knolls, Bay of Plenty, New Zealand. Remarks: Awhiowhio osheai differs from the two known Habitat: Encrusting on basalt rocks. Depth range species of Herengeria in the lack of acanthose microxeas 159–318 m. that are abundant on the ostial face of H. vasiformis and H. auriculata, the lack of hair-like diactines, and the Description: simpler ornamentation of the desmas. The ectosomal Morphology conical to cup-shaped with thick straight micropolyrhabds are also simpler, with no branches walls (Fig. 13A,C). NIWA18333 is a vertical plate with and only small irregular protrusions. In A. osheai the a spherical outline (Fig. 13B); the base is broad and oscules are clearly visible and raised on the concave elliptical, margins of the plate are smooth. surface of the cup. A. osheai is so far known only from Dimensions 33–70 mm wide, 22–53 mm high, 8–12 seamounts in the Bay of Plenty, New Zealand. mm thick, base 18–46 mm long, 13–26 mm wide. Texture stony. Key diagnostic characters: Surface — outer surface of cup smooth to undulating • Relatively regular bean- or rod-shaped micropoly- with a granular texture; oscules, c. 0.1 mm diameter, rhabds on the surface. scattered regularly over the concave face in cup-shaped • No surface microxeas and diactines. specimens, flush with the surface in some specimens • Thick straight-walled cup with a smooth dimpled and raised in others; ostia not visible. concave surface; can easily be confused in the field Colour dull yellow (3B3) in life and in ethanol. with Aciculites pulchra but the oscules on the latter Choanosomal skeleton a dense reticulation of desmas are more pronounced and flatter and the sponge is and massive intricate zygoses that are pierced by cir- a curvaceous ear rather than straight-walled. cular channels (Fig. 13D). Ectosomal skeleton a dense regular crust of micropo- n. sp. (Fig. 14) lyrhabds over the dichotriaene cladomes. Awhiowhio unda Megascleres (Table 12) desmas are relatively slender Material examined: with 2-4 clones, heavily nodulose with single or mul- Type material: Holotype NIWA18337 from NIWA tiple nodular spires (Fig. 13D). Zygoses are massive, Stn K826. highly intricate and spread laterally along the clones in mature sections of the sponge skeleton, forming a dense Type locality: Southern South Fiji Basin. aggregation with trabeculae. Desmas have a granular brown core, immature desmas are sparsely tuberculate Distribution: Southern South Fiji Basin. with relatively smooth thin clones; dichotriaenes (Fig. 13D–F) have a relatively long rhabd and short robust Habitat: Attached to a piece of dead coral. Depth range clads, with frequent trichotriaene modifications. 390–490 m. Microscleres (Table 12) relatively regular rod-shaped acanthose micropolyrhabds (Fig. 13G), with bulbous Description: ends and a central expansion, straight or curved or Morphology comprising a highly foliose thick-walled in a few cases, abruptly angled centrally. Streptasters lamella attached along the base, the lamella with mul- (Table 12) with extremely fine long rays are common tiple infoldings and undulations, the margin slightly in the choanosome (Fig. 13G). expanded and flared (Fig. 14A). Dimensions 64 mm long, 45 mm wide, 35 mm high, Etymology: Named for Dr Steve O’Shea, formerly of lamella 7–10 mm thick. NIWA Wellington, for his considerable knowledge of Texture stony. Table 12. Spicule dimensions (µm) of Awhiowhio osheai n. sp.

Clone (thickness) Cladome Micropolyrhabd (L) Streptaster (L) Overall size (Max ø) (Thickness) of arch Rhabd (L)

NIWA18332 50–60 206 (150–250) 27 (20–38) 30 (25–38) (holotype) c. 200–500 461 (325–700) 4–5

NIWA18333 40–50 183 (100–240) 24 (15–30) 30 (23–38) (paratype) c. 200–500 490 (260–675) 2–5

41 Figure 14. Awhiowhio unda n. sp.

42 Table 13. Spicule dimensions (µm) of Awhiowhio unda n. sp.

Clone (thickness) Cladome (Max ø) Microrhabds (L) Streptaster (L) Overall size of arch Rhabd (L) (Thickness) NIWA18337 20–50 195 (150–240) 18 (15–23) 25 (20–33) (holotype) c. 200–300 296 (150–420) 5–8

Surface smooth, granular. Type locality: Graveyard Seamount, Chatham Rise. Colour coffee brown (5F6) in preservative. Choanosomal skeleton comparatively delicate, formed Distribution: Known only from the Graveyard by small, relatively slender tripod-shaped dicrano- seamount complex, a series of small knolls and hills clones. Small streptasters are abundant in the choano- on the Chatham Rise. some amongst the desmas (Fig. 14B). Ectosomal skeleton a shallow layer of robust micro- Habitat: Silty hard substratum. Depth range 757– scleres. Dichotriaene cladomes line the ectosome (Fig. 875 m. 14B). Description: Megascleres (Table 13) — desmas small, frequently Morphology comprising a large erect plate with a broad tripod-shaped and relatively slender, the clones are undulation in the plane of the upper margin, the mar- smooth with smooth nodules (Fig. 14C). The nodules gin indented by what appear to be large shallow pits on the apex of tripod desmas are spires with bumpy c. 5–7 mm wide. Base of sponge thick, flanged (Fig. tubercles (Fig. 14B). Zygosis is mostly terminal and 15A). relatively weak as articulation is via saddle-shaped Dimensions c. 300 mm wide, c. 210 mm high, base clasps to adjacent clones. Dichotriaenes are small and 130 wide, 40 mm wide; thickness of plate at base 44 the cladomes are short and robust (Fig. 14B, D). mm, at middle of plate 28 mm, margin 12 mm. Microscleres (Table 13) — choanosomal microscleres Texture brittle, stony, the margin slightly flexible. are streptasters verging on spirasters with sharply Surface macerated, the underlying skeleton smooth, pointed rays (Fig. 14E). Ectosomal microscleres are indicating its possible condition in life. irregular thick micropolyrhabds that may have an ir- Colour dirty white in macerated condition. regular spiraster-shape (Fig. 14D,E). Choanosomal skeleton of heavily zygosed desmas Etymology: Named for the wave-like morphology of that form ‘bridges’ (Fig. 15C,D). Only a single triaene the sponge walls and the microscleres (unda, L.) was seen (Fig. 15E). Megascleres (Table 14) — large desmas that form Remarks: Awhiowhio unda is similar to A. osheai in that large rounded arches or ‘bridges’ joined laterally via it has small, relatively smooth desmas with nodulose saddle-shaped zygomes (Fig. 15C–E). Nodular spires clones, but the streptasters and micropolyrhabds are arise from the arched side of the primary clone. Desmas more spiraster-like. The species has a unique wavy are large and thick, with a granular centre. Clones are morphology. smooth with occasional nodules that are ‘sculpted’ around the termini. Smaller, more intricately branched Key diagnostic characters: desmas (Fig. 15 B) were observed in preparations of the • Frilled wave-like sponge wall. abraided surface of the sponge, where short clones arise • Small non-tuberculate desmas compared to other on all sides of the primary linear clone. Dichotriaenes species. with robust straight conical rhabd and clads. Microscleres absent owing to macerated condition Awhiowhio sepulchrum n. sp. (Fig. 15) of sponge.

Material examined: Etymology: Named for the type locality, the Graveyard Type material: Holotype NIWA18338 from NIWA Stn Seamount on the Chatham Rise (sepulchrum, Latin, TAN0104/002. grave). Table 14. Spicule dimensions (µm) of Awhiowhio sepulchrum n. sp.

Clone (thickness) Cladome (Max ø) Overall size of arch Rhabd (L) Overall size of smaller ramose desmas NIWA18338 70–150 375–900 (holotype) c. 900–1200 450–750 c. (600–800) – (300–400)

43 Figure 15. Awhiowhio sepulchrum n. sp.

44 Remarks: The specimen described above is tentatively whole. In doing so, it became obvious that Callipelta, assigned to Awhiowhio n. gen. because the absence Daedalopelta, and Homophymia differ considerably from of diagnostic microscleres precludes full identifica- typical corallistid genera like Corallistes, Herengeria, and tion. The specimen was dead before collection, with Neoschrammeniella in having pseudotriaenes, smooth nothing remaining of the surface and mesohyl or all monocrepid pseudotetraclones, and amphiasters as microscleres and non-desma megascleres, with the microscleres. Callipelta, Daedalopelta and Homophymia exception of a single dichotriaene, having sloughed were accordingly transferred to the Neopeltidae. off the desma skeleton. Pisera and Lévi (2002g) upheld this decision in their The assignment to Awhiowhio is based on the mor- recent review of the Neopeltidae but they disagreed phology of the desmas. These are relatively smooth with the hypothesis of Kelly (2000) that Neopeltidae dicranoclones with a few nodulose spires on the outer and Corallistidae are closely linked in the suborder Di- edge of the arched tripod-shaped primary clone. The cranocladina, a subgroup of lithistid sponges that have basic shape and ornamentation of the desma, with monaxial desmas. Recent rDNA data analyses (Kelly nodules rather than multituberculate fungiform pro- & McCormack unpubl.) indicate that the Neopeltidae jections, separates the sponge from Herengeria, but the and Corallistidae are indeed linked and that this group strong lateral bridge-forming zygoses strongly resem- is quite separate from tetracladine, rhizomorine, and bles that of both Awhiowhio and Herengeria, linking it megamorine taxa. to Corallistidae. Despite the lack of diagnostic microscleres and ac- Homophymia Vacelet & Vasseur, 1971 cessory megascleres, the assignment of a species name to NIWA18338 is justified by the possession of several Fan-shaped or clavate Neopeltidae with monocrepid unique features. NIWA18338 is the only known rep- pseudotetraclones, smooth monocrepid pseudophyl- resentative of the family Corallistidae in the Chatham lotriaenes as ectosomal megascleres, and a single form Rise region. With the exception of Neoschrammeniella of amphiaster microsclere. antarctica from the Ross Sea, Corallistidae are common in waters only as far south as Raukumara Plain off East Type species: Homophymia lamellosa Vacelet & Vasseur, Cape. The species is also readily distinguished from 1971. other Corallistidae by the greater size and simpler ornamentation of the dicranoclone desmas. A more Homophymia stipitata Kelly, 2000 (Fig. 16A–F) detailed evaluation of the genus assignment is not possible without living specimens in which megasclere Homophymia stipitata Kelly, 2000: 9, figs 5–8. and microsclere complements are intact. Material examined: Key diagnostic characters: NIWA Stations: I96 [NIWA holotype H-705 • Sponge forms a very large flat plate. (NIWA7503), NIWA paratypes P-1152 (NIWA4775) • Very large arched simple dicranoclones with a and NIWA P-1155 (NIWA4778)]; J953 [NIWA paratype dense brown granular core. P-1153 (NIWA4776, BMNH 1995.3.30.2)]; J954 [NIWA paratype P-1156 (NIWA4779, BMNH 1995.3.30.5), Family NEOPELTIDAE Sollas, 1888 NIWA25611]; P8 (NIWA25645); P10 [NIWA paratype P-1154 (NIWA4777)]; Z9747 [BMNH 2005.9.23.1 (frag- Neopeltidae Sollas, 1888: 344; Topsent 1894: 290; Schrammen ment of 0CDN6731-Z)]. 1903: 20; Kelly 2000: 281 (partim). NMNZ Stations: NMNZ Por. 615. Daedalopeltidae de Laubenfels, 1936: 184. Distribution: Wanganella Bank (holotype); West Nor- Thickly encrusting, planar, pedunculate, fungiform folk Ridge; west of Pandora Bank; western continental lithistid demosponges with monaxial pseudodiscotri- slope of Northland, New Zealand. aenes, pseudophyllotriaenes, and/or siliceous discs; desmas are monocrepid with a complex branching Habitat: Collected from coralline and shell rubble shape resembling tetraclones (pseudotetraclones); banks on the northeastern continental shelf of North- may be accompanied by strongyles; microscleres are land. The sponge is typically attached to hard substrata amphiasters and/or acanthose microxeas. but may be partially buried. This species is part of a biodiverse invertebrate assemblage in which hydroids, Remarks: The discovery of a second species of Homo- soft corals, bryozoans, and ascidians are typical. Other phymia from continental-shelf banks off northeastern lithistid sponges include Aciculites pulchra, Discodermia New Zealand led Kelly (2000) to redefine the genus proliferans, and several species of Neoschrammeniella and and to evaluate the integrity of the Corallistidae as a Pleroma (Kelly 2000). Depth range 192–757 m.

45 Figure 16. Homophymia stipitata Kelly, 2000.

46 Description (after Kelly 2000): waters; it is spherical with an apical depression and a Morphology parsnip-shaped (stipitate) with an irregular short stalk. A fourth species has since been collected globular body that bulges in places (Fig. 16A); apex is from the New Caledonian Norfolk Ridge seamounts depressed, forming a translucent oscular sieve through (Kelly unpubl.), the morphology of which is similar to which exhalant canals are clearly visible (Fig. 16B). that of H. stipitata but with a smaller, penicillate body Dimensions of body 40–50 mm diameter, about one and only a slight expansion of the apical region. The third of total length of 150–220 mm, elevated upon and typical form of the type species, H. lamellosa, is easily tapering to a relatively long fine stalk ranging from distinguishable from the above three species in that it 20 mm diameter at the junction of the body, to 5 mm is a folded lamellate fan and is apparently restricted diameter at the base of the stalk. to the Western Indian Ocean. Texture barely compressible, stalk is rigid but rela- It is noteworthy that very similar ‘phyllotriaenes’ tively flexible along entire length. to those of H. stipitata were figured from the Oamaru Surface smooth but slightly undulating and irregu- Diatomite (c. 35 Ma) (Hinde & Holmes 1892, pl. 14, lar, granular to the touch. Exhalant canals are visible fig. 1), and described from the Tutuiri Greensand of on surface of body and stalk beneath the translucent Chatham Island (c. 63–50 Ma). Superficially, the Tu- ectosome (Fig. 16B). tuiri, Oamaru, and Recent spicules appear to be very Colour in life ivory (4B3), in preservative light similar, and were indeed considered to be congeneric golden brown (5B7), apical depression dark grayish by Kelly and Buckeridge (2005). Closer examination of blue (20E4). the Tutuiri specimens, however, indicates that while Choanosomal skeleton composed of huge smooth they are indeed identical to the Oamaru material, they monocrepid pseudotetraclones (Fig. 16C) and tracts of differ from those of Recent specimens in the possession large oxeas that radiate through the stalk towards the of a triaenose inception canal, which is clearly illus- apical surface. Amphiasters are abundant throughout trated in the Hinde and Holmes (1892) figure. the choanosome and the ectosomal region. Ectosomal skeleton packed with pseudophyllotri- Key diagnostic characters: aenes, the sinuous ‘fingers’ of the ‘cladome’ partially zygose with neighbouring ‘cladomes’ and desmas • Parsnip-shaped with a grey-blue apical depres- below the surface. The ‘rhabds’ of these pseudophyl- sion. lotriaenes are perpendicular to the surface. • Surface covered with pseudotriaenes with an Megascleres (Table 15) — desmas of the choanosome extremely sinuous cladome and short irregular are huge smooth ramose monocrepid pseudotetra- rhabd. clones (Fig. 16C). Ectosomal megascleres are pseu- • Desmas are very big and resemble tetraclones but dophyllotraienes (Fig. 16D,E) in which the ‘rhabd’ is the epirhabd is only a short bar. irregular in shape and thickness, is sinuous, and the • One type of amphiaster. cladome is polyfurcate with sinuous branches. The spicules are laminated and granular in appearance. Callipelta Sollas, 1888 Diactines are strongyloxeas, curved, syringe-like, in two size classes. Pillow-shaped, spherical, clavate, and multiclavate Microscleres (Table 15 Ms 1) — amphiasters (Fig. Neopeltidae, with relatively small irregular mono- 16F) with curved spines on the distal two thirds of crepid desmas that resemble tetraclones; ectosomal oxeote rays. megascleres are moncrepid pseudodiscotriaenes to pseudophyllotriaenes with strongly incised denticulate Remarks: Homophymia stipitata was the second species margins on the ‘clads’, the upper surface of which may to be described, the genus being previously known be tuberculate; microscleres are amphiasters. only from Madagascar. Schlacher-Hoenlinger et al. (2005) recently described a third species of Homophy- Type species: Callipelta ornata Sollas, 1888. mia from Kaimon Maru Seamount in New Caledonian

Table 15. Spicule dimensions (µm) of Homophymia stipitata Kelly, 2000.

Desma (Clone L/T) Cladome (Max ø) Diactines 1 (L) Ms 1 (L) Desma (Overall size) Rhabd (L) Diactines 2 (L) Epirhabd (L) NIWA7503 147–588/20–59 627 (465–711) 1055 (809–1372) 15 (12–19) (holotype) 400–682 254 (186–323) 472 (308–617) 193 (147–225)

47 Callipelta punctata Lévi & Lévi, 1983 (Fig. 17) rigid (Fig.17C); ectosomal megascleres are pseudophyllotriaenes with a short rhabd that is often slightly Callipelta punctata Lévi & Lévi, 1983: 118, pl. 6, fig. 1, pl. 8, obliquely set (Fig. 17B). The centre of each rhabd has a figs 1–3, 9. crepis about 30 µm long as in the desmas. The cladome Material examined: is not precisely perpendicular to the rhabd, and often appears slightly concave in profile. The pseudoclads NIWA Stations: KAH0011/30 (NIWA18339); may be composed of a pseudoprotoclad, which may TAN0413/117 (NIWA24045). split to form two branches, or may remain single. The end of the clad may branch again; the whole is very Distribution: New Caledonia (holotype); Rungapapa delicate and lacy in appearance. The upper surface Knoll, Bay of Plenty. of the cladome is denticulate (Fig. 17B). Diactines are relatively short fusiform oxeas. Habitat: Numerous small specimens encrusting speci- Microscleres (Table 16) amphiasters with long fine mens of Erylus nigra Bergquist, 1968 (Porifera, Astro- acanthose rays (Fig. 17D). phorida, Geodiidae) and volcanic basalt substrata, with numerous other invertebrates including immature Remarks: These sponges are most similar to Callipelta Reidispongia coerulea. Depth range 159–360 m. punctata Lévi & Lévi, 1983 from New Caledonia in Description: terms of spicule morphology and dimensions. In particular, the upper surfaces of the pseudophyllotriaenes Morphology semi-spherical to columnar, with a single have tuberculate projections, a character that appears small oscule on the slightly depressed apex of the to be unique to C. punctata and species of Macandrewia sponge (Fig. 17A). NIWA24045 from TAN0413/117 is (Macandrewiidae). a tiny slightly raised encrustation. The New Zealand sponges are more columnar than Dimensions 3–7 mm diameter, 5–10 mm high, oscule those described from New Caledonia, but the punctate 0.5 mm diameter. The New Caledonian material was surface is characteristic. In gross morphology they are reported as 15–18 mm diameter and 10 mm high. perhaps more similar to Callipelta sollasi Lévi & Lévi, Texture stony, crumbly. 1989 from about 100 m depth off Luzon and Mindoro Surface microscopically rough and punctate, slightly Islands in the Philippines, but this latter species has hispid with projecting diactines; occasional aquiferous pseudophyllotriaenes with a smooth cladome surface, canals converging on the oscules incising the surface which is also much wider (300–350 µm) than those of or appear below it. C. punctata. Colour in life cream (4A3) and in ethanol golden The only other described species, C. cavernicola brown (5D7). (Vacelet & Vasseur, 1965), C. thoosa Lévi, 1965, and C. Choanosomal skeleton composed of monocrepid mixta Vacelet, Vasseur & Lévi, 1976 are quite different desmas and short diactines. morphologically and in terms of spiculation, and all Ectosomal skeleton lined with the scale-like cladome three species are known only from submarine tunnels of the pseudophyllotriaenes. and grottos on the coast of Madagascar. Callipelta mixta Megascleres (Table 16) are monocrepid pseudotetra- is probably a species of Neopelta as it contains discotri- clone desmas with a short epirhabd in which a short aene-like megascleres and acanthose microxea. slender bar crepis (30 µm) resides (Fig. 17C). Two thick Despite extensive collection, this species has so far clones arise from each end of the epirhabd; these have been found only at a single New Zealand location, the a granular core, the terminus of each clone tuberculate Rumbles Seamount in the Bay of Plenty. or spinose, the zygosis between spicules relatively

Table 16. Spicule dimensions (µm) of Callipelta punctata Lévi & Lévi, 1988.

Desma (Clone L/T) Pseudophyllotriane Diactines (L) Amphiaster (L) Desma (Overall size) cladome (Max ø) Rhabd (L)

New Caledonia (holotype) 60–80/20–30 200–300 350–500 20 ― 75–150 New Zealand NIWA18339 80–130/30 224 (150–280) 400–600 16 (15–18) 180–230 100 (50–140)

48 Figure 17. Callipelta punctata Lévi & Lévi, 1983.

49 Key diagnostic characters: Distribution: Volcano L, Kermadec Ridge, northeastern • Tiny columnar or encrusting sponges (< 20 mm New Zealand. diameter) with a tiny apical oscule and a rough Habitat: Encrusting hard basalt substrate. Depth range punctuate surface. 154–240 m. • Desmas and pseudophyllotriaenes are granular in the centre. Description: • Desmas have a single fine bar crepis (monocre- Morphology pillow- or cushion-shaped, edges thinning pidial) in the epirhabd. to the substratum. • Can be mistaken for Macandrewia spinifoliata in the Dimensions 10 mm long, 7 mm wide, 4 mm thick. field; the presence of amphiasters and the lack of Texture stony, brittle. ectosomal smooth strongylote oxeas (present in Surface smooth. Macandrewia spinifoliata) in Callipelta punctata, are Colour in ethanol honey yellow (5D6). indicative. Choanosomal skeleton composed of small robust monocrepidial desmas that form very intricate zygoses. Neopelta Schmidt, 1880 Amphiasters are abundant in the choanosome. Ectosomal skeleton partially lost, but in patches pseu- Pillow-shaped and columnar Neopeltidae with small dodiscotriaenes form a scaled surface. complex tuberculate monocrepid desmas that resem- Megascleres (Table 17) — monocrepid pseudotetra- ble tetraclones; ectosomal megascleres are moncrepid clone desmas (Fig. 18B) with a short thick epirhabd circular, oval to elongate, sculpted and angular and with four short thick clones that extend into very tuber- ridged discs; some are pseudodiscotriaenes with an culate termini. The clones are less tuberculate towards oblique pseudorhabd projection that contains the the smooth epirhabd. On the sponge surface, clones monaxial crepis; other megascleres may include slen- are palmate and similar to moose antlers in shape. The der curved oxea-like diactines that occur singly or zygoses are very complex and highly robust. Ectos- in bundles; microscleres are amphiasters; acanthose omal megascleres (Table 17) are pseudodiscotriaenes microxea occur in some species. with a flat to slightly concave disc, which are circular, elongate, or multilobate in overall shape, with circular Type species: Neopelta perfecta Schmidt, 1880. indentations on the edge of the disc, usually one per Remarks: Neopelta plinthosellina Lévi & Lévi, 1988 from spicule. The upper surface of the discs is ornamented New Caledonia differs from the type species N. perfecta (Fig. 18A), some with swirls of small evenly spaced in that it lacks acanthose microxea and has elongate tubercles, some with what appear to be knobbed ridges and highly sculpted discs. The diagnosis above has leading to the centre of the disc, and some which lack been broadened to accommodate the peculiar sculpted either embellishment. discs of N. plinthosellina, and the lack of microxeas and Microscleres (Table 17) are amphiasters with long choanosomal oxeas and tuberculate regular palmate fine rays. desmas in both southwestern Pacific species. Etymology: Named for the pillow- or cushion-shaped gross morphology (pulvinus, L.). Neopelta pulvinus n. sp. (Fig. 18) Remarks: This single specimen of Neopelta pulvinus Material examined: was initially identified as a species of Callipelta (Ne- Type material: Holotype NIWA18340 from NIWA Stn opeltidae) because it has ectosomal monocrepidial TAN0205/75 spicule slide labelled ‘NIWA18340’. pseudodiscotriaenes, amphiasters as microscleres, and lacks the characteristic acanthose microxea of Type locality: Volcano L, Kermadec Ridge, northeast- the type species (Neopelta perfecta Schmidt, 1880) and ern New Zealand. N. imperfecta Schmidt, 1880, both from the Caribbean.

Table 17. Spicule dimensions (µm) of Neopelta pulvinus n. sp.

Desma (Clone L/T) Pseudodiscotriaene disc (Max ø) Amphiaster (L) Desma (Overall size) Pseudorhabd (L) Desma crepis (L) Epirhabd (L) New Zealand NIWA18340 100–250/50–70 227 (88–300) by 140 (63–220) 19 (15–25) (holotype) 300–500 20 30 18

50 Figure 18. Neopelta pulvinus n. sp.

51 The ectosomal discotriaenes are, however, quite differ- for Lepidospongia Dendy, 1924, which was preoccupied. ent from those of most species of Callipelta in that they He transferred Lepidothenea incrustans to the Kaliap- are discate rather than phyllamentous. The discotriaene sidae de Laubenfels (= Phymaraphiniidae in part) pseudorhabd in this new species is also unlike those of owing to the possession of what he perceived to be the pseudophyllotriaenes of Callipelta species, which ‘phyllotriaenes’ instead of dichotriaenes (de Laubenfels are almost always well defined and perpendicular 1936). Bergquist (1968) did not re-examine this mate- to the cladome. In N. pulvinus the rhabds are solid, rial but retained it in the original family Theonellidae, short, frequently with two flanges that adhere to the no doubt convinced that the report of what appeared underside of the disc, and the pseudorhabd is often to be rhabds on some discs, i.e. ‘discotriaenes’, abun- positioned obliquely or perpendicular to the under- dant ectosomal microrhabds, and vestigial style-like side of the disc. This unusual rhabd morphology has megascleres, indicated broad affinity with species in been described for both species of Caribbean Neopelta the genus Discodermia. Pisera (pers. comm. 2004) has (Sollas 1888), albeit rarely, but not for the only other suggests that while the sponge does indeed resemble known species, New Caledonian N. plinthosellina Lévi a juvenile Discodermia, albeit without desmas, it also & Lévi, 1988. resembles members of the genus Neopelta. It is his opin- Neopelta plinthosellina, like N. pulvinus n. sp., lacks ion that Lepidothenea is probably a younger synonym acanthose microxeas and both species have very similar of Neopelta. desmas; some are typically pseudotetraclones with smooth or tuberculate clones and heavily tuberculate Lepidothenea incrustans (Dendy, 1924) (Fig. 19) zygoses, but some are plamate and spreading as illustrated by Lévi and Lévi (1988, pl. 6). The desmas Lepidospongia incrustans Dendy, 1924: 317, pl. 13, figs 1–3. of the southwestern Pacific species also differ from Lepidothenea incrustans: de Laubenfels 1936: 176; Bergquist those of the Caribbean species, which are complex with 1968: 64. smooth rays and are frequently branched (see Pisera & Lévi 2002g). It is thus with some confidence that Material examined: None. the Kermadec Ridge specimen is assigned to the same group as N. plinthosellina from New Caledonia, but little Distribution: Holotype Terra Nova Station 90, approxi- confidence is placed in the assignment of both species mately 20 km southwest of the summit of Great King to the Caribbean genus Neopelta. Despite the obvious Island (34º24’S, 172º06’E) (Harmer & Lillie 1914). differences between the Caribbean and southwestern Habitat: Dendy (1924) described the habitat of ‘Terra Pacific species ofNeopelta , additional specimens of both Nova’ Station 90 as being remarkable in that it yielded species, particularly of N. pulvinus, are required for full thirty-eight species of sponges. The depth was about characterisation of these differences, and separation 183 m and the substratum was rocky. The sponge from Neopelta as a new genus. formed a thin crust on a small block of basalt with numerous other encrusting organisms. Key diagnostic characters: Description: • Tiny encrusting cushion sponge. • Surface has scale-like discs with tiny projecting Morphology a thinly spreading encrustation. The tubercles on the upper surface and usually one or specimen was described as partially rubbed off the more circular indentations. substratum. • Desmas are frequently palmate. Dimensions 10 5 10 mm, the thickness was not given by Dendy (1924). Texture crisp, delicate. Incertae sedis Surface formed of overlapping scales, interrupted by fairly numerous delicate minute conical apertures (Dendy Lepidothenea de Laubenfels, 1936 1924), presumably oscules. Sponge forms a very thin encrustation, without des- Colour in life described as ‘beautifully’ silvery. The mas; skeleton comprises an upper and lower adherent sandwiched choanosome was dark brown and dry. of overlapping scale-like discs or occasionally discotri- Skeleton comprised of an upper and lower layer of over- aenes, between which is a thin choanosome. Microscle- lapping discate scales, the latter adherent to the underlying res are microrhabds. (Modified from Dendy 1924). substratum, separated by an equally thin choanosome. Some thin vestigial megascleres were found to be associated with Type species: Lepidospongia incrustans Dendy, 1924. the rhabds of some discotriaenes. Megascleres almost exclusively irregular oval to circu- Remarks: As pointed out by Bergquist (1968), De lar discs, perfectly smooth and structureless, the margins Laubenfels (1936) proposed the new name Lepidothenea occasionally indented, without any trace of a shaft or axial

52 Figure 19. Lepidothenea incrustans (Dendy, 1924) canals; a few have a stout, well-developed rhabd, and some lack of amphiasters, and the lack of microxeas common cladomes have a reticulate pattern; maximum diameter of in the family. It is recommended that Lepidothenea be cladome 250 µm (Fig. 19A, B). retained as a valid genus, though incertae sedis, within Microscleres are microrhabds, fusiform, straight or the Neopeltidae. centroangulate, slightly roughened; c. 20 µm long and 2 Key diagnostic characters: µm thick (Fig. 19C). • Tiny encrustation. Remarks: The genus Lepidothenea de Laubenfels is • Surface has scale-like discs. known from a single very small specimen dredged • The only other spicules are acanthose micro- from 183 m off the Three Kings Islands. No further rhabds. material is available and the sponge has not been recollected since. Family MACANDREWIIDAE Schrammen, The phylogenetic affinity ofLepidothenea incrustans 1924 remains in contention. Some characters (occasional ‘discotriaenes’ with a stout ‘rhabd’, microrhabds, rare Macandrewiidae Schrammen, 1924: 39. vestigial styles) weakly link this species with Discoder- mia (Theonellidae) and (microrhabds, monocrepidial Polymorphic lithistid demosponges with dentate phyl- discs) Neopelta (Neopeltidae) but the sponge lacks lotriaenes, smooth ectosomal strongylar oxeas, and desmas. The lack of desmas in a lithistid sponge has a desmas with a triaenose crepis. (Modified from Pisera precedent, however; a species of Theonella from Van- & Lévi 2002m). uatu is known to lack ectosomal phyllotriaenes and desmas; all that remain is the strongyle megasclere Macandrewia Gray, 1859 skeleton and microrhabds in the ectosome (Kelly unpubl.). On balance, L. incrustans has some affinity with Cup-shaped, massive-irregular, branching, or lamel- species in the Family Neopeltidae, but differs in the late Macandrewiidae with dentate phyllotriaenes, unique thinly encrusting form, the lack of desmas, the the clads of which form intricate swirling antler-like

53 patterns, desmas with a triaenose crepis resembling with differential development of some clads, the edges tetraclones overall; the ectosome is encrusted with crenulate and indented. smooth strongylar curved oxeas. Microscleres may Microscleres (Table 18) smooth curved spicules with include roughened centrotylote microxeas. strongylote ends, the ends (Fig. 20D).

Type species: Macandrewia azorica Gray, 1859. Remarks: The details of spiculation conform to the holotype description except that the desmas are triae- Macandrewia spinifoliata Lévi & Lévi, 1983 neose, not monocrepidial rhizoclones as described by (Fig. 20) Lévi and Lévi (1983). While Pisera and Lévi (2002m) refuted the hypothesis put forward by Kelly (2000) Macandrewia spinifoliata Lévi & Lévi, 1983: 116, fig. 8, pl. 6, that this genus has affinities with the Neopeltidae, it fig. 3, pl. 8, figs 4–6. is interesting to note that the phyllotriaenes of Macan- drewia spinifoliata are very similar in structure and size Material examined: to the desmas, as in the neopeltid genera Homophymia NIWA Stations: KAH9907/48 (spicule slide labelled and Callipelta. The contention that the phyllotriaenes ‘MKB 1829’); I372 (NIWA25780). of Macandrewia are pseudophyllotriaenes is supported by the fact that the short axial filaments of the crepis do Distribution: New Caledonia (holotype); Doubtless not extend in the plane of the cladome to give rise to Bay, North Cape; west of White Island, Bay of Plenty, monoclads or deuteroclads as in the triaenes in sponges New Zealand. with tetraclone desmas. The phylogenetic affinities of this monogeneric family remain elusive. Habitat: Encrusting basalt rock. Depth range 211– A single specimen of Macandrewia spinifoliata was 395 m. recognised during close inspection of the minute spe- Description: cies that grow on basalt boulders dredged from the Bay of Plenty seamounts. These boulders have proved to be Morphology nubbin-shaped; Lévi and Lévi (1983) re- a rich source of new species of lithistid sponges. corded numerous specimens that were digitate and ramified with knobby ends. Key diagnostic characters: Dimensions 13 mm wide, 4 mm high. • Tiny encrustation (on basalt rock on Bay of Plenty Texture fragile, crumbles easily. seamounts). Surface smooth. • Surface has feather-like triaenes and curved stron- Colour in ethanol ivory (4B3). gylote oxeas. Choanosomal skeleton composed of small pseu- • Microscleres in the shape of asters are absent. dotetraclone desmas that form a close mesh and intricate zygoses. Ectosomal skeleton lined with the phyllotriaene Family PLEROMIDAE Sollas, 1888 cladomes, encrusted with strongylote oxeas and rough Megamorina Zittel, 1878: 99 (partim). microrhabds. Pleromidae Sollas, 1888: 312; Topsent 1894: 290; Wilson 1925: Megascleres (Table 18) comprising small robust 461; de Laubenfels 1955: E50. desmas (Fig. 20A) with a triaenose crepis (one actine is Pleromatidae: Lendenfeld, 1903: 140. longer than the other three) that resemble tetraclones Pléromides: Moret, 1926: 118. (Fig. 20B). The clones are smooth around the epirhabd but can be tuberculate towards the zygoses, which are Massive, conical, plate- to ear-shaped or globose lithis- highly intricate. Phyllotriaenes (Fig. 20C) with straight tid demosponges; surface usually granular or hispid conical rhabd, cladome irregularly circular to polyclad, with projecting megascleres, texture stony. Desmas smooth or nodulose monocrepid megaclones with Table 18. Spicule dimensions (µm) of Macandrewia spinifoliata Lévi & Lévi, 1983.

Desma (Clone L/T) Phyllotriaene disc (Max ø) Diactines (L) Microxea (L) Desma (Overall size) Rhabd (L)

New Caledonia Lévi & Lévi, 1983 ― 200–300 400 60–70 (holotype) 150–250 100–125

New Zealand ‘MKB 1829’ 120–150/30–34 200 (150–250) x 272 (160–450) 378 (250–425) 80 (63–108) 200–250 121 (90–150)

54 Figure 20. Macandrewia spinifoliata Lévi & Lévi, 1983.

55 saddle- or cup-shaped zygomes; ectosomal spicules feld 1903: 140; de Laubenfels 1955: E50, fig. 31.3; Lévi & are dichotriaenes and/or anatriaene-like megascleres; Lévi 1983: 105, fig. 2, pl. 2, figs 5–8; Pisera & Lévi 2002d: microscleres are spirasters, amphiasters, streptasters 324–326, figs 5–7; Kelly 2003: 116, figs 2A,B, 3. in various sizes, and finely spined microxeas and/or Material examined: lightly acanthose style-like monactines. (From Kelly 2003). NIWA Stations: I92 (BMNH 2002.3.4.1, NIWA25061); KAH9907/50 (NIWA18344); X140 (NIWA18341, Remarks: Two valid extant genera, Pleroma and Ana- NIWA18342); Z2092 (NIWA18343); Z9400 derma, are recognised in Pleromidae, with four species (NIWA25620). known from seamounts and banks in the Southwest IRD Stations: LITHIST CP9 (NIWA18519, NIWA18520), Pacific including New Zealand and New Caledonia DW7 (NIWA18521). (Pisera & Lévi 2002d; Kelly 2003; Kelly et al. 2003). Distribution: Makutu coast, Fiji Islands (holotype); Havannah, south coast of New Caledonia, and Éponge

Pleroma Sollas, 1888 Seamount, south New Caledonian slope of Norfolk Lyidium Schmidt, 1870: 84. Ridge; Lord Howe Seamount Chain; Norfolk Ridge Pleroma Sollas, 1888: 312; Lendenfeld 1903: 140; de Laubenfels south of Norfolk Island; Challenger Plateau, northern 1955: E50; Pisera & Lévi 2002d: 324, figs 5–7. slopes; west of White Island, Bay of Plenty.

Massive irregular encrustations, thick folded plates, or Habitat: The sponges were attached to hard rocky globular with a flattened apex; the surface granular and substrata on the flanks of seamounts. Depth range hispid in places, texture stony. Desmas are smooth or 230–860 m. fully nodulose monocrepidial megaclones with cup- shaped, saddle-shaped, or irregular zygomes with Description (modified from Kelly 2003): indented margins, megascleres are large oxeas and Morphology a curved ear-like semicircular plate, at- dichotriaenes, microscleres are roughened microxeas tached to the substratum along the base of the sponge and one to two types of microspined spiraster, amphi- (Fig. 21A), or with a restricted base of attachment (Fig. aster, and streptaster. (From Kelly 2003). 21B). The edges of the plate are rounded in profile. Dimensions of body up to 240 mm high and wide, Remarks: Hinde and Holmes (1892) illustrated several 10–30 mm thick. megamorine microfossil desmas from the Oamaru Texture stony. Diatomite (c. 35 million years ago), at Oamaru in North Surface velvety on the upper concave surface, Otago. These are remarkably similar to desmas of granular on the lower convex surface, oscules are not Pleroma turbinatum (Hinde & Holmes 1892, pl. 13, fig. visible (see Fig. 21A). 25) and P. menoui (Ibid., pl. 13, figs 26, 27). Colour in life beige, in preservative cream to beige. Type species: Pleroma turbinatum Sollas, 1888. Choanosomal skeleton composed of megaclones with saddle-shaped zygomes. Abundant microxeas fill the Pleroma turbinatum Sollas, 1888 (Figs 21, 22) interstices between the desmas (Fig. 22A). Ectosomal skeleton an encrustation of microscleres Pleroma turbinatum Sollas, 1888: 312, pl. 33, figs 5–7; Lenden- and dichotriaenes.

Table 19. Spicule dimensions (µm) of Pleroma turbinatum Sollas, 1888 (summarised from Kelly 2003).

Desma (Clone L/T) Cladome (Max ø) Diactines (L) MO (L) Ms1 Ms2 Desma (Overall size) Rhabd (L)

Fiji Sollas (1888) ―/50–90 448 1350 180–210 24–30 ― (holotype) 500–800 960

New Caledonia Lévi & Lévi 200–300/70–90 360–590 1500–1700 130–150 12 ― (1983) 500–550 1000–2000

New Zealand Kelly (2003) 100–800/60–100 375–750 1000–2000 130–165 10–20 12 375–625 650–875

56 Figure 21. A, B, Pleroma turbinatum Sollas, 1888; C, P. menoui Lévi & Lévi, 1983; D, P. aotea Kelly, 2003.

57 Megascleres (Table 19) — choanosomal desmas are IRD Stations: LITHIST CP9 (IRDR1827; NIWA18522, smooth megaclones with a light brown granular core NIWA18523); DW11 (NIWA18524). and a monoaxial crepis, with 3–6 clones emanating from both sides of a relatively long epirhabd (Fig. Distribution: South of New Caledonia (holotype), 22A,E). The zygomes are oval saddle- shaped expan- Éponge and Kaimon Maru Seamounts, south New sions (Fig. 22E). Long fine oxeas extend through the Caledonian slope of Norfolk Ridge; western continental ectosome. Dichotriaenes have very short protoclads slope of Northland; west of Three Kings Ridge; Colville and long deuteroclads. Kelly (2003) noted that the Ridge; southern South Fiji Basin; Wanganella Bank, distal end of the rhabd is rounded in New Zealand northwestern New Zealand; Raukumara Plain, north- specimens. ern Bay of Plenty; Volcano E, Kermadec Ridge; West Microscleres in two forms, a regular microspined Cavalli, South Cavalli, and Cavalli Seamounts; west of spiraster ((Table 19, Ms 1; Fig. 22I) and a smaller mi- White Island and Tuatoru Knoll, Bay of Plenty. crospined spiraster/streptaster (Table 19, Ms 2; Fig. 22K) with rays of uneven length and orientation. These Habitat: Attached to hard substrata on continental shelf latter microscleres, in some specimens, have very thick banks. Found with a range of other lithistid sponges, rays. Microxeas are roughened and fusiform, straight molluscs, bryozoans, brachiopods, echinoderms, hy- and centrally thickened. droids, and stylasterid hydrocorals (Kelly 2000). Depth range 179–1131 m. Remarks: No additional material has been collected since the original description of New Zealand speci- Description (modified from Kelly 2003): mens in Kelly (2003 Morphology a highly irregular multilobate encrusting plate, occasionally tubular (Fig. 21C) or a thick fan Key diagnostic characters: composed of fused lobes. Immature sponges are thick • Very thick, stony, ear-shaped. flattened ear-shaped plates (see Kelly 2003). • Easily be confused with Herengeria vasiformis or Dimensions of most mature specimens are c. 21 cm species of Neoschrammeniella but the smooth desmas long, 11 cm high, 14 cm wide. are characteristic. Texture stony. Surface extremely hispid with long oxeas protruding Pleroma menoui Lévi & Lévi, 1983 (Figs 21, 22) beyond 10 mm (Fig. 21C). Colour in life beige, in preservative cream to Pleroma menoui Lévi & Lévi, 1983: 103; Kelly 2003: 119, figs beige. 2C,D, 4, 6. Choanosomal skeleton composed of megaclone desmas (Fig. 22B,C) with cup- shaped zygomes, sitting on Material examined: on top of the other (Fig. 22C). Extremely long oxeas NIWA Stations: J953 (NIWA18347, NIWA25606, protrude more than 10 mm from the sponge surface BMNH 1995.3.30.1); S562 (NIWA18345); S572 (Fig. 22B). (NIWA18346); K826 (NIWA18350); K872 (NIWA18349); Ectosomal skeleton charged with strongylospirasters, K A H 0 2 0 4 / 0 9 (NIWA18355, NIWA18356); microxeas, dichotriaenes, and long hair-like diactines KAH0204/27 (NIWA18353); KAH0204/30 (Fig. 22B). (NIWA18354); KAH0204/44 (NIWA18357); Megascleres (Table 20) — megaclones are smooth T A N 0 2 0 5 / 2 0 (NIWA18351); TAN0205/60 with a laminated appearance and a pale brown granu- (NIWA18352); TAN0413/085 (NIWA25054); lar core containing the monaxial crepis; 2–5 clones ema- TAN0413/093 (NIWA25053); X152 (NIWA18348); nate from both sides of a thick epirhabd, but typically Z9263 (NIWA25059). radiate in one direction (Fig. 22B,C,F) forming dipods, NMNZ Stations: NMNZ Por. 580; NMNZ Por. 591. Table 20. Spicule dimensions (µm) of Pleroma menoui Lévi & Lévi, 1983 (summarised from Kelly 2003).

Desma (Clone L/T) Cladome (Max ø) Diactines (L) MO (L) Ms1 Ms2 Desma (Overall size) Rhabd (L)

New Caledonia Lévi & Lévi 400/120–180 1200 12000 170–225 15–18 10 (1983) 300–700 700–1400

New Zealand Kelly (2003) 375–700/125–200 600–1250 12000 170–250 12–20 17 575–1250 1125–2325

58 Figure 22. A, E, I, K, Pleroma turbinatum Sollas, 1888; B, C, F, H, L, P. menoui Lévi & Lévi, 1983; D, G, J, P. aotea Kelly, 2003. 59 tripods, and quadripods (see also Lévi & Lévi 1983, pl. west of White Island, and Whakatane Seamount, Bay 2, figs 1–2). The epirhabd is slightly thicker than the of Plenty. clones. The zygomes are spherical and cup-shaped. Diactines are long and fine. Dichotriaenes are large, Habitat: Attached to hard substrata on seamounts. slender and curved, the deuteroclads and protoclads Depth range 230–1538 m. are quite long and fine (Fig. 22B). Microscleres in two forms, a regular strongylos- Description (modified from Kelly 2003): piraster derived from a spiraster with rays encased in Morphology of immature sponges comprisinge spheri- additional silica (Table 20 Ms 1; Fig. 22H), and a smaller cal knobs with a restricted base of attachment. As streptaster (Table 20 Ms 2; Fig. 22L). Roughened micro- the sponge matures it flattens and the apical surface xeas are thick and slightly angulate centrally. becomes slightly concave with one or more oscular depressions. Immature sponges usually have a single Remarks: Specimens of Pleroma menoui Lévi & Lévi apical oscule and slightly depressed vestibule beneath are identical in New Zealand and New Caledonian this oscule, whereas the largest have numerous oscules specimens; the desmas are very large, arched, smooth spread across the surface (Fig. 21D). and thick, with cup-shaped zygomes, and microscleres Dimensions of mature specimens can reach up to include characteristic strongylospirasters. 80 cm diameter by 35 mm high, but the majority of Since the revision and discussion by Kelly (2003), specimens in the collection average 40 mm diameter the species range has been extended to the Kermadec by 30 mm high. Ridge and Cavalli Seamounts of northeastern New Texture stony. Zealand. Surface smooth and granular. Colour in life cream to beige (4A3). Key diagnostic characters: Choanosomal skeleton composed of tripod-like nodu- • Stalked or convoluted plate sponge with thick walls, lose megaclones (Fig. 22D). Centrotylote microxeas stony and very hairy. are abundant in the outer choanosome and ectosome • Desmas with distinctive smooth clones in the form along with 2 forms of microscleres. Large oxeas may of a tripod, dipod, or quadripod, ends of clones have protrude a short distant from the surface. sucker-like zygomes. Ectosomal skeleton with abundant centrotylote mi- • Microscleres include acanthose bean-like spicules. croxeas and two forms of microscleres line the surface. Dichotriaenes are also abundant in the ectosome. Pleroma aotea Kelly, 2003 (Figs 21, 22) Megascleres (Table 21) — megaclones are irregular with 2–4 clones emanating from a short, slightly bul- Pleroma aotea Kelly, 2003: 123, figs 2E–I, 5, 6. bous epirhabd (Fig. 22D, G) containing a monaxial crepis. The surface of the desma has relatively well- Material examined: defined low mounds or nodules. The clones emanating NIWA Stations: AUT 137; I97 [NIWA holotype H-807 from the epirhabd are often shorter and thinner than (NIWA7613)], [NIWA paratype P-1276 (NIWA5097, the epirhabd and are frequently dichotomous. The BMNH 2002.3.4.2)]; KAH0204/08 (NIWA18358); zygomes are enormous with an oval profile and they KAH0204/44 (NIWA18359, NIWA18360, NIWA18361); are occasionally dichotomous with irregular indented KAH9907/50 (NIWA18362); S568 [NIWA paratype margins (Fig. 22G). Dichotriaenes are robust with a P-1278 (NIWA5101)]; TAN0413/016 (NIWA25052); broad cladome, the protoclads are thick and straight, U591 [NIWA paratype P-1277 (NIWA5098)]; Z9262 and the deuteroclads are sturdy and often split at the (NIWA25607). extreme end (Fig. 22D). Large oxea are common and protrude a short distance from the surface. Distribution: Challenger Plateau, northern slope; Microscleres are amphiaster/streptasters in two Wanganella Bank, West Norfolk Ridge (holotype); Tui size categories (Table 21, Ms 1, 2; Fig. 22J) with very Seamount, Three Kings Ridge; western flank of Three fine sharply pointed rays. Roughened microxeas are Kings Ridge; Cavalli and South Cavalli Seamounts; thick, straight, and cigar-shaped (see Fig. 22D) and often centrotylote. Table 21. Spicule dimensions (µm) of Pleroma aotea Kelly, 2003 (summarised from Kelly 2003).

Desma (Clone L/T) Cladome (Max ø) Diactines (L) MO (L) Ms1 Ms2 Desma (Overall size) Rhabd (L)

New Zealand Kelly (2003) 375 (275–500)/125–225902 (725–1125) 2000–3000 130–160 20–22 10 703 (475–950) – 1086 (825–1325) 1096 (700–1375)

60 Remarks: Pleroma aotea is relatively easy to recognise they have lateral notch-like zygoses; the desmas of in the field as it is mushroom- shaped and very stony. the outer choanosome are like sinuous oxeas. Other The desmas are very characteristic of the species and megascleres include long sinuous projecting diactines, differ considerably in both size and shape from those plagiotriaenes, and/or dichotriaenes. Choanosomal mi- of both P. turbinatum and P. menoui. croscleres are microxeas, amphiasters, or streptasters; Kelly et al. (2003) reported the discovery of numer- ectosomal microscleres are micropolyrhabds. ous fossil sponges from the Late Eocene–Early Oli- gocene Ototara Limestone at Kakanui, North Otago, Type species: Costifer vasiformis Wilson, 1925. that were considered to be indistinguishable from P. aotea at the gross morphological level. Because the Costifer wilsoni Lévi, 1993 (Figs 23, 24) sponges are now solid calcite, a full confirmation of this species identity is not possible. However, desmas very Costifer wilsoni Lévi, 1993: 20, fig. 5B, pl. 2, fig. 2, pl. 9, figs similar to those of P. aotea were also reported from the 1–2. Tutuiri Greensand (Late Paleocene) of Chatham Island Material examined (Kelly & Buckeridge 2005), confirming a southern dis- : tribution in New Zealand during this time. NIWA Stations: KAH0204/08 (NIWA18364, NIWA18365, NIWA18366); S572 (NIWA18363); Z9262 Key diagnostic characters: (NIWA25617, NIWA25618). • Small mushroom-like sponges with a restricted base NMNZ Stations: NMNZ Por. 597, NMNZ Por. 712. and slightly concave bumpy surface. IRD Stations: LITHIST CP9 (NIWA18525). • Desmas are highly characteristic with the nodulose surface and saddle-shaped zygomes. Distribution: South New Caledonian slope (holotype), Stn CP105, and Éponge Seamount, south New Caledo- Family ISORAPHINIIDAE Schrammen, nian slope of the Norfolk Ridge; Challenger Plateau, northern slope; Wanganella Bank, West Norfolk Ridge; 1910 West of Three Kings Ridge; Cavalli Seamount, north- Helomorinidae Schrammen, 1910: 128. eastern New Zealand. Isoraphiniidae Schrammen, 1924a: 38; de Laubenfels 1955b: E51. Habitat: Attached to hard rocky substrata in seamount environments. IRD Stn CP9 was extremely diverse Lithistid demosponges with smooth, linear oxea-like with sea fans and numerous Herengeria vasiformis, Neo- sinuous heloclones bearing lateral notch-like zygoses, aulaxinia spp., Geodia spp., and Pleroma menoui (Kelly plagiotriaenes, and/or dichotriaenes, and accessory unpubl.). Depth range 206–904 m. diactines. Microscleres are acanthose microxeas, amphiaster/strepasters, and micropolyrhabds (modified Description: from Pisera & Lévi 2002l). Morphology a thick, heavy ear-shaped or semicircular lamella attached to the substratum by a short stem REMARKS: The Isoraphiniidae is based on two species 30–40 mm wide and c. 10 mm high that projects from of the Recent genus Costifer —the type species C. vasi- the back of the sponge base (Fig. 23A); margin of la- formis Wilson, 1925 from the Philippines and C. wilsoni mella curved. Lévi 1993 from New Caledonia (Pisera & Lévi 2002l). Dimensions 400 mm wide, 160 mm high, 40 mm The family is represented by numerous Mesozoic thick (IRD Stn CP9). New Zealand specimens 70–200 genera. Several new specimens of C. wilsoni from New mm wide, 40–130 mm high, 20–25 mm thick. Zealand and one from New Caledonia, described here, Texture tough but slightly flexible, not stony. add to knowledge of this previously rare genus. Surface generally smooth but furry or granular to the touch. Colour in life peach-coloured (6A6), in preservative Costifer Wilson, 1925 dark mustard-brown (5E8). Costifer Wilson, 1925: 461. Choanosomal skeleton composed of large thick heloclones that are abundant throughout the choanosome, Vase-shaped, lamellar, or auricular lithistid demos- arranged randomly and sometimes radially at the ponges; surface hispid, texture incompressible, slightly surface (Fig. 23B). Heavily notched desmas appear flexible. Choanosomal desmas are heloclones in two to be commoner in the sponge interior, while the ox- categories — those of the deep choanosome have ir- eote desmas are commoner on the exterior projecting regular expansions and restrictions along the spicule, through the ectosome. Microxeas are abundant and the ends are bulbous strongylote or attenuated, and pack the choanosome.

61 Ectosomal skeleton packed with micropolyrhabds. measure as they are broken, but appear on some specimens Megascleres (Table 22) — heloclone desmas in two to be greater than 10 mm long. morphologies: (1) outer choanosomal desmas are long, Microscleres — microxeas, very finely roughened smooth, sinuous, and oxea-shaped, attenuated at the with a small central swelling, slightly curved or angled tips, frequently bent over at one end (Table 22 D1, from midpoint of spicule (Fig. 23B, 24A,D); micropolyr- Figs 23B,E, 24A,B); (2) deeper choanosome desmas habds (Table 22, Ms 1, Fig. 24D,E) acanthose, irregular, are shorter irregularly polytylote with strongylote, with bulbous projections, curved or linear or stepped truncate or bulbous attenuated ends and lateral sad- in length. Some are irregularly spiraster- shaped. dle-shaped notches (Table 22 D2; Fig. 23C,D, 24C). Acanthose amphiaster/streptasters (Table 22, Ms 2, Some desmas have a hollow axial canal extending Fig. 24E) with fine rays. the full length of the spicule, and others have a fine axial crepis that ends in a series of small branches in Remarks: There is some minor variation between the some spicules. Ectosomal triaenes are plagiotriaenes spicule dimensions of the New Caledonian holotype or dichotriaenes in equal numbers (Fig. 23E). The pro- material and the New Zealand material but the New torhabd of the dichotriaene cladome is much longer Zealand specimens conform to the holotype descrip- than the two deuteroclads on each of the three clads that tion in all other aspects. Further to Lévi’s (1993) de- make up the cladome. Plagiotriaenes have a much broader scription, the ratio of plagiotriaenes to dichotriaenes cladome than dichotriaenes, and shorter rhabds. Long fine can vary from specimen to specimen; in NIWA18366, diactines project from the surface; these are impossible to plagiotriaenes are more abundant, while in NMNZ Por.

Figure 23. Costifer wilsoni Lévi, 1993.

62 Figure 24. Costifer wilsoni Lévi, 1993.

63 Table 22. Comparison of the spicule dimensions (µm) of Costifer wilsoni Lévi, 1993 with the type species C. vasiformis Wilson, 1925 (P cladome = plagiotriaene dimensions, D cladome = dichotriaene dimensions).

D 1 (L) D 2 (L) D Cladome (Max ø) Microxeas Ms 1 Ms 2 (W) (W) Rhabd (L) (L) (L) (L) P Cladome (Max ø) Rhabd (L)

New Caledonia Lévi (1993) 1500–4000 ― 600–1000 130–220 30 12 (holotype) 40–120 ― 800–1000

New Zealand NMNZ Por. 597 2488 (2000–3000) 1907 (1550–2550) 1468 (1250–1650) 211 (175–232) 24 15 100 (70–150) 152 (100–200) 813 (650–1000) 1750 1125 NIWA18366 2438 (1650–2700) 2469 (2375–2600) 1375–1625 183 (100–220) 29 15 200–225 100–125 2375 1937 (1500–2250) 1775 (1625–1700)

Philippines C. vasiformis 1880–2370 ― 525–830 90–165 18–25 15 (Wilson, 1925) 220–280 850–1310

597 dichotriaenes dominate. Furthermore, the desmas Microfossil heloclones indistinguishable from those are in two recognisable categories — the desmas of the of extant Cosfiter wilsoni were found in the Tutuiri outer choanosome are predominantly long, smooth, Greensand (Teurian–basal Waipawan) in the Chatham sinuous, and oxea-shaped, and frequently bent over Islands (Buckeridge & Kelly 2005), confirming a more at one end, while the deeper choanosomal desmas southern distribution for this genus during the Paleo- are generally shorter and more irregular with uneven gene. expansions and contractions along the axis and with strongylote, truncate or bulbous ends. Along the axis Key diagnostic characters: of these latter spicules are conspicuous lateral saddle- • Large, heavy, ear-shaped; may be mistaken for shaped notches indicating the proximity of adjacent Pleroma turbinatum but the former is flexible while spicules. Desmas may have a hollow axial canal extend- the latter is stony. ing the full length of the spicule, or a fine axial crepis. • Presence of heloclones in choanosome. Either form of desma may have a hollow or thread-like axial morphology. Family Scleritodermidae Sollas, 1888 Pisera and Lévi (2002l) suggested that C. wilsoni is very similar to the genus holotype Costifer vasiformis Scleritodermidae Sollas, 1888: 346; Wilson 1925: 463; de Wilson, 1925 and that it differs only in the possession Laubenfels 1936: 165. of dichotriaenes and in the absence of lateral notches in Massive-amorphous, encrusting, flabellate, ear-, cup- or the heloclones. Examination of this new material has vase-shaped. Desmas are thorny or tuberculate rhizo- afforded a clearer differentiation (see Table 22) between clones; ectosomal spicules when present are irregular the two species; the microxeas, heloclones and plagi- curved smooth or acanthose spicules with strongylote otriaenes of C. vasiformis are considerably smaller than ends, some are polyrhabdose, or larger acanthose tylos- those of C. wilsoni, and the morphology of the former trongyles; microscleres when present are sigmaspires. is vase-shaped with a rough irregular surface. This (Modified from Pisera and Lévi 2002b). contrasts considerably with C. wilsoni, which is smooth and ear-shaped. Note that lateral notches are certainly Remarks: A partial rhizoclone desma skeleton, very present in New Caledonian material (NIWA18525); similar to those in extant species of Scleritodermidae, contrary to what Pisera and Lévi (2002l) suggested, was found in the Tutuiri Greensand (Teurian–basal these desmas with notches are perhaps commoner in Waipawan) in the Chatham Islands (Buckeridge & the deeper choanosome. Kelly 2005), confirming a more southern distribution for this family during the Palaeogene.

64 Microscleroderma Kirkpatrick, 1903 form slightly concave circular lamellae attached to the substratum at the base (Fig. 25B). Microscleroderma Kirkpatrick, 1903: 173. Dimensions 45 mm high, 60 mm wide and 30 mm Taprobane Dendy, 1905: 103; Dendy 1922: 7; Wilson 1925: thick, lamella thickness 5–7 mm. 468. Texture stony, but easily snapped. Surface smooth, velvety on both surfaces, but more Cup- or ear-shaped with a hirsute, smooth, or tuber- on the concave surface. culate concave surface; desmas are thorny rhizoclones; Colour ivory (4B3) in life and in ethanol. choanosome also may contain long oxeote diactines; Choanosomal skeleton a reticulation of very spiny ectosomal megascleres absent; microscleres are sigma- rhizoclone desmas (Fig. 25C). spires. (Modified from Pisera & Lévi 2002b). Ectosomal skeleton with fine diactines projecting beyond both surfaces, diactines are longer on the mar- Type species: Microscleroderma hirsutum Kirkpatrick, gin of the lamella. C-shaped, S-shaped, and spiralled 1903. sigmaspires are abundant in the outer 500 µm of the ectosome and are less common in the deep choano- Microscleroderma novaezelandiae n. sp. some. (Fig. 25; Table 23) Megascleres (Table 23)—rhizoclone desmas (Fig. 25C), simple, arched, with numerous elongate single or bifud spines arising from the arch, usually from Material examined: the outward curved side. The lower extremities of the Type material: Holotype NIWA18367 from NIWA Stn arch may divide into 2–3 short clones that are involved Z7190; paratype NIWA18368 from NIWA Stn Z7187. with zygosis (Fig. 25D, upper right). The appearance of ladder-like “fibres” is given by zygosis of the clones Distribution: Unfortunately, no information is available with adjacent desmas at each end of the spicule. These on the collection localities for these two specimens, align in parts of the skeleton to form what appear as other than that they were collected in New Zealand the longitudinal beams of a ladder, the arch or “beam” waters. Collection details are limited to those in Ap- forming the rungs. Long oxeote diactines in two size pendix 3. categories, the longer and thicker confined to the margin of the lamella, and the shorter and finer confined Habitat: Attached to rocky or coral substrata. Depth to the concave surface. These latter diactines have fine range unknown. fusiform whip-like ends. Microscleres (Table 23) comprise spined S-shaped Description: and C-shaped sigmaspires (Fig. 25C,D). Morphology comprising shallow curved circular lamellae joined to form a tubular flared vase-like structure; Remarks: Several characters differentiate Microsclero- margin of lamellae smooth, rounded, more hirsute derma novaezelandiae from the Indo-PacificM. herdmani than the flat surfaces (Fig. 25A). Immature specimens (Dendy, 1905) and New Caledonian M. stoneae Lévi

Table 23. Comparison of the spicule dimensions (µm) of Microscleroderma novaezelandiae n. sp., M. stonae Lévi & Lévi, 1983, and M. herdmani (Dendy, 1905).

Desma 1 (L)/ Clone (Thickness) Diactines 1 (L) Diactines 2 (L) Sigmaspires (L) Desma 2 (L)/ Clone (Thickness) M. stoneae New Caledonia 300/30 900-1000/5-13 ― 7-10 150/5

M. novaezelandiae NIWA 18367 300/30 <2500/15 532(300-775)/1-2 13(10-15) (holotype) ―

M. herdmani Philippines 200-300 2-6 mm ― 6-8 (Wilson 1925) ―

New Caledonia 200/30 900-1500 ― 10-12 (Lévi & Lévi 1983) ―

65 Figure 25. Microscleroderma novaezelandiae n. sp.

66 & Lévi, 1983. Microscleroderma forms a curved folded Description: lamella that is spherical in general outline; M. herdmani Morphology a thick plate attached to the substratum by a is vase-shaped and the lamellae of mature specimens broad ridge; upper surface verrucose with meandering roll inward along the vertical plane to form a complex flattened ridges and nodules (Fig. 26A), with regularly of vases. Both surfaces of M. novaezelandiae are smooth scattered oscules that have slightly raised edges; lower and velvety, unlike the concave surfaces of M. herdmani surface is ridged and flatter than upper surface. and M. stoneae, which are hirsute and have mamillate Dimensions 60 mm by 30 mm wide, 20 mm high in projections (Dendy 1905; Wilson 1925; Lévi & Lévi middle of specimen (including basal ridge), 10–12 mm 1983). The sigmaspires are larger in M. novaezelandiae thick at margin. than those described for M. herdmani by Dendy (1905) Texture stony. and in New Caledonian material (Lévi & Lévi 1983) Surface granular, with flattened ridges and nod- (see Table 23), and it has two size categories of diac- ules. tines, with the thicker longer diactines on the margin Colour dark mustard brown (5E8) in life and in of the sponge. preservative. While M. novaezelandiae is more similar to M. sto- Choanosomal skeleton a dense reticulation of desmas, neae Lévi & Lévi, 1983, which forms a simple shallow being more cavernous on the external surfaces. bowl, careful surface scrapes did not reveal the highly Ectosomal skeleton dark brown and membranous and ramified superficial desmas that typify this New Cal- contains a relatively thick oblique to tangential layer edonian species. of acanthorhabds (Fig. 26B). These microscleres are also present within the outer choanosome but become Key diagnostic characters: less common deeper within. Sigmaspires are present • Small thin-walled cup or plate. in the outer choanosome and ectosome, but these are • S- or C-shaped sigmaspires and thin diactines in not very common. two size categories. Megascleres (Table 24) thorny rhizoclone desmas, elongate with short, branched termini. Scleritoderma Sollas, 1888 Microscleres acanthorhabds (Table 24; Fig. 26C), slightly centrally thickened, with strongylote or faintly Scleritoderma Sollas, 1888: 315. tylote ends, straight, curved or faintly S-shaped; sigmaspires C- or S-shaped (Fig. 26D). Massive, flabellate, encrusting, or plate-like, with nodes or are cup-shaped; desmas are spinose rhizo- Remarks: The single New Zealand specimen conforms clones; ectosomal spicules are faintly S- or C-shaped reasonably closely to the type species Scleritoderma acanthorhabds with strongylote or faintly tylote ends, flabelliformis Sollas, 1888, although the morphology is microscleres are sigmaspires. (Modified from Pisera slightly different (not strictly flabelliform). Sollas (1888) and Lévi 2002b). mentioned that “the sponge is sometimes in its most irregular and incurved form, attached by the convex Type species: Scleritoderma flabelliformis Sollas, 1888. side of the folded plate.” This latter morphology better describes the New Zealand specimen. Other species of Scleritoderma flabelliformis Sollas, 1888 Scleritoderma include S. nodosum Thiele, 1900 from Ter- (Fig. 26) nate and Tulear, Madagascar, but this has deep channels and columns that appear as nodes on the surface Scleritoderma flabelliformis Sollas, 1888: 316, pl. 35, figs 26–50; of the sponge and is quite different from the plate-like Lévi & Lévi 1989: 47, fig. 15, pl. 2, fig. 3. morphology of S. flabelliformis. Scleritoderma camusi Lévi & Lévi, 1983 from New Caledonia forms a deep-blue Material examined: shallow cup sponge of regular morphology. NIWA Station: S572 (NIWA18369). The rare exotyles mentioned by Lévi and Lévi (1989) were absent in the New Zealand specimen and Distribution: Kei Islands, Indonesia (holotype); Philip- the holotype. These Philippine specimens differ sig- pines; west of Three Kings Ridge, New Zealand. nificantly from the holotype morphology; the figured specimen is heavily digitate with rounded nodular tips Habitat: Attached to hard substrata. An undescribed (Lévi & Lévi 1989, pl. 2, fig. 3), extending knowledge species of Hamacantha (Poecilosclerida, Hamacanthi- of the range of morphologies in this species, if indeed dae) encrusts the surface in small patches. Depth range it is S. flabelliformis. 403–530 m.

67 Figure 26. Scleritoderma flabelliformis Sollas, 1888.

68 Table 24. Spicule dimensions (µm) of Scleritoderma flabelliformis Sollas, 1888.

Desmas (L) Acanthorhabd (L) Exotyles (L) Sigmaspires (L) Clone (Thickness) (Thickness)

Indonesia (Sollas 1888) 240–500 80–90 ― 10 (holotype) 39 13

Philippines (Lévi & Lévi 1989) ― 60–95 160 (rare) 8–10 5–7 2

New Zealand NIWA18369 200–400 75 (60–90) ― 9–10 30–45 5–7

The rare occurrence of S. flabelliformis in New Zea- (NIWA18390); TAN0413/109 (NIWA18386); land probably indicates the southernmost limit of this TAN0413/111 (NIWA18391); TAN0413/118 species, which is commoner in the Indo-Pacific. (NIWA18387, NIWA18388); TAN0413/170 (NIWA18385); TAN0413/173 (NIWA18392); Key diagnostic characters: TAN0413/174 (NIWA18393); TAN9915/3B • Massive plate with a verrucose (corrugated and (NIWA25528); V473 (NIWA18373); X128 (NIWA18374); nodular) upper surface. Z8487 (NIWA18371); Z9041 [NIWA18376, NIWA25033]; • Presence of curved, weakly S- or C-shaped acan- Z9742 [NIWA18823 (0CDN6692-J)]; Z9747 (NIWA25060 thorhabds with faintly bulbous (tylote) ends, and (fragment of 0CDN6730-Y]); Z9890 (NIWA25616); sigmaspires. Z9892 (NIWA25615). NMNZ Stations: NMNZ Por. 308; NMNZ Por. 562; Aciculites Schmidt, 1879 NMNZ Por. 563; NMNZ Por. 564; NMNZ Por. 570; NMNZ Por. 571; NMNZ Por. 573; NMNZ Por. 620. Aciculites Schmidt, 1879: 29. Distribution: West Norfolk Ridge; southern Three Kings Cushion-shaped, with raised or tubular oscules, Ridge; western continental slope, Northland; west of encrusting, or massive-columnar with blind digits; Pandora Bank; North Cape; Louisville Seamount choanosomal desmas are rhizoclones with spinose Chain; Kermadec Ridge; Cavalli Seamount; Ngunguru zygomes; ectosomal megascleres are curved strongyles Bay, Northland; outer Bay of Islands; off Tutukaka; to tylostrongyles with one or both ends acanthose, Rungapapa, Tumokemoke, Mahina, and Tuatoru arranged tangentially or paratangentially in the ecto- Knolls, west of White Island, and Mayor Island, Bay some; microscleres absent. (Modified from Pisera and of Plenty; east of Gisborne; south of East Cape; Mahia Lévi 2002b). Peninsula.

Type species: Aciculites higginsi Schmidt, 1879. Habitat: CRRF specimens were dredged from ‘coarse sand and rubble’, and ‘rocky reef substrate’. The base Aciculites pulchra Dendy, 1924 (Fig. 27, 28) of the sponge encrusts hard substrata. Depth range 80–1050 m. Aciculites pulchra Dendy, 1924: 315, pl. 6, fig.1–1a; Bergquist 1968: 63, fig. 30, pl. 10a. Description: Morphology vase-shaped with foliose lamellae (Fig. Material examined: 27A,B), or ear-shaped with a flat lamella (Fig. 27C, D). NIWA Stations: E636 (NIWA18377); F993 Dimensions 50–200 mm wide, 10–15 mm thick, up (NIWA18372); I14 (NIWA18370); J954 (NIWA18375, to 30 mm at the base of some specimens. NIWA25609); KAH0011/30 (NIWA18378); Texture stony. KAH0011/40 (NIWA18379, NIWA18380); Surfaces sharply differentiated; the exhalent surface, KAH0011/41 (NIWA18381); KAH9301 (NIWA25610); usually the concave surface, is smooth and crowded KAH9907/48 (NIWA18382, NIWA18383, NIWA18384); with numerous regularly dispersed low hemispherical P8 (NIWA25608, NIWA25646); RAPUHIA2-15 oscular projections c. 0.5 mm high; the other (inhalent) (NIWA25779); TAN0413/068 (NIWA25040); surface is smooth, granular, and irregular growth rings TAN0413/092 (NIWA18389); TAN0413/099 may be visible.

69 Figure 27. Aciculites pulchra Dendy, 1924.

Colour in life — reddish-brown ex- Table 25. Spicule dimensions (µm) of Aciculites pulchra Dendy, 1924. terior (9E6) and pinkish interior (9B4); in preservative light brown. Desmas (L) Tylostrongyles (L) Diactines (L) Choanosomal skeleton a dense re- Clone (Thickness) Thickness ticulation of rhizoclone desmas (Fig. New Zealand 28B); these are less complex in the Dendy (1924) 340 340 ― outer choanosome of both surfaces. (holotype) ― 12 Ectosomal skeleton on the inhalent surface consists of radially arranged Bergquist (1968) 361 (300–387) 368 (314–411) 566 (510–663) 10 (6–13) 23-30 tylostrongyles that arise from a region of maturing desmas (Fig. 28A). On NIWA18371 210–300 285 (200–400) ― the exhalent surface these megascle- 20–30 7–13 res gradually become obliquely and then tangentially layered around the NIWA18372 300–400 440 (300–580) ― curved margin of the lamella where 20–30 5–10 tylostrongyles form a thick tangential layer over the exhalent surface.

70 Figure 28. Aciculites pulchra Dendy, 1924.

Megascleres — thorny rhizoclone desmas (Table Mahia Peninsula. These oxeas were described as being 25; Fig. 28A–C) with long crepi, terminating in 1- to radially disposed and arranged singly or in groups of 3-spined cladi, spines single or bifurcate; zygosis is up to four just below the oscular surface. Occasional lateral and terminal and may involve the spinose pro- spicules fitting this description were found in a new jections (Fig. 28C); tylostrongyles (Fig. 28D,E) (‘rhabdi’ spicule preparation of Bergquist’s specimen. However, of Dendy 1924: 316), curved or straight with slightly large ‘true’ oxea were not described from Dendy’s holo- tylote acanthose proximal head and smooth strongy- type and have not been found in any of the specimens lote distal terminus. Most megascleres are strongylote examined in this study. with one end acanthose. There is considerable variation in the dimensions Microscleres absent. and morphology of the tylostrongyles; in Bergquist’s Mahia specimen, the tylostrongyles are markedly ty- Remarks: Bergquist (1968) found large stout ‘oxeas’ in lote, almost bulbous at the proximal end, and faintly addition to the typical tylostrongyles in specimens off tylote hammer-like at the distal end. In some specimens

71 oxeote diactines have been found but they are rare and Habitat: Coarse sandy rubble. Depth range 198– probably derived from the tylostrongyle megasclere; 319 m. they are syringe-like with a thick hollow inception channel, and one or both ends are slightly sinuous and Description: elongate-fusiform. Some also have faintly tylote ends. Morphology tree-like with one or more thick, sometimes These “diactines” are no longer than the tylostrongyles. hollow columns fused along the stem or above where Some specimens have tylostrongyles that taper distally the digits emerge; stem circular in profile or flanged if to an oxeote point and it may be these that were seen more than one stem is present (Fig. 29A). Some speci- in Bergquist’s specimen. mens present a more columnar shape (Fig. 29B). The The spiculation of this species is thus quite variable. stem flares and gives rise to numerous branches that However, the vase-like morphology with raised oscules, may be hollow at the base, each diverging to form one coupled with the presence of a range of tylostrongyles or more smaller branches. These terminate in a rounded (dimensions and morphology) is diagnostic. or pointed blind tip (Fig. 29C). The overall impression Aciculites pulchra Dendy is almost identical morpho- is of an irregular squat tree with an irregularly truncate logically to the Indo-PacificA. ciliata Wilson, 1925 with top that is compressed in one plane. Oscules are not reddish-brown colouration, a vase-like foliose or ear- apparent, although there are soft areas at the tips of the shaped lamella, and raised oscular projections on the sponge that may function as exhalant areas. exhalant surface. Examination of numerous specimens Dimensions of the holotype are 100 mm wide at the from the Philippines and Papua New Guinea (courtesy apex, 50 mm thick, 70 mm tall, and 50 mm wide at the of CRRF) reveal that the tylostrongyles in A. ciliata are base. NMNZ Por. 583 is 85 mm wide at the apex, 60 mm much longer (up to 540 µm long) and are irregularly thick, 110 mm tall, and 30 mm wide at the base. Sec- bulbous at one end, however. The spicules also taper ondary branches are c. 15 mm thick, smaller branches to a fine blunt distal point. This species also possesses c. 5–10 mm thick. much longer fine tylostrongyles that resemble oxeas, Texture of the stem and lower branches is stony, tips up to 850 µm long; these are radially arranged along are fragile and soft. the inhalant surface. The outer choanosome in A. ciliata Surface granular, rough, encrusted by numerous Wilson is highly cavernous with elongate immature epizoites including other sponges, bryozoans, and desma. tubeworms (Fig. 29A). Colour in life dark grayish-blue (20E4), in preser- Key diagnostic characters: vative light golden brown (5B7). Tips of branches are • Vase- or ear-shaped lamella with differentiated paler than the digits of the apex. surfaces. Choanosomal skeleton composed of thorny rhizoclone • Reddish-brown colouration. desmas that are less complex and dense in the outer • Presence of tylostrongyles with one end rough- choanosome. ened. Ectosomal skeleton composed of tylostrongyles ar- • Could be mistaken for Neoschrammeniella fulvodes- ranged obliquely to paratangentially over most of the mus but that species does not have raised oscules. body surface and radially at the growing tips of the sponge. Aciculites manawatawhi n. sp. (Fig. 29) Megascleres (Table 26) rhizoclone desmas are elongate and arched with short clads at the ends of the desma (Fig. 29D). Ornamented with short thick Material examined: bifurcate spines; overall a thick, robust spicule. Zygosis Type material: Holotype BMNH 2005.9.15.1 from appears to be predominantly terminal. Tylostrongyles NIWA Station Z9747 [fragment of 0CDN6740-L]; para- or ‘oxytyles’ with pronounced acanthose bulbous type NIWA18394 from NIWA Stn J953. heads and strongylote or tylote acanthose distal ends NIWA Station: P14 (NIWA25623). in some spicules the proximal end is bent sharply as NMNZ Station: NMNZ Por. 583. in a rhabdostyle (Fig. 29E). Microscleres absent. Type locality: West of Pandora Bank, northwestern New Zealand. Etymology: Named for the largest island in the Three Kings group, Manawa Tawhi (Great Island), the type Distribution: West Norfolk Ridge; west of Pandora locality. Manawa tawhi (New Zealand Māori) means Bank, and Middlesex Bank, northwest of Three Kings ‘heart food’ (manawa, heart, breath, emotions), but it Islands; western continental slope, northeastern New may mean ‘the heart beckons’ (tawhi, food, beckons) Zealand. (Weno Iti & Guy Penny pers. comm.).

72 Figure 29. Aciculites manawatawhi n. sp.

73 Table 26. Spicule dimensions (µm) of Aciculites mana- and tylostrongyles protrude ‘head-first’ through the watawhi n. sp. surface, an architecture that is well described by Pisera and Levi (2002f) and Muricy and Minervino (2000)], Desmas (L) Oxytyles (L) which do not form a tangential layer, and the species Clone Thickness typically forms club-shaped columns with a deep broad (Thickness) cavity. Pisera and Levi (2002f) noted that Gastrophanella New Zealand also has long smooth styles and long oxeas, absent BMNH 2005.9.15.1 200–400 380 (300–500) from A. manawatawhi and A. oxytylota. The diagnosis (holotype) 30–40 8–10 of Aciculites has been expanded to include species with NIWA18394 300–400 345 (280–430) oxytylotes in a paratangential layer. 50 8–12 It is clear that the dimensions of tylostrongyles and 51 desmas cannot be used as a species-level differentia- tor in Aciculites as the lengths of these spicules in the NMNZ Por. 583 250–300 390 (340–500) different species are very similar, ranging from about 20–40 8–10 200–500µm, and the full range often present in some species. (A notable exception is A. spinosa, which has quite small tylostrongyles ranging from 125 to 275 Remarks: Aciculites manawatawhi is unique in the genus µm.) The shape of the tylostrongyle appears to be more in terms of gross morphology and is easily differenti- diagnostic; A. oxytylota and A. manawatawhi have ox- ated from other Indo- and Southwest Pacific species, ytyles with pronounced bulbous heads, but the latter which are ear- or cup-shaped (A. pulchra Dendy, 1924 species has tylote bulbs on both ends of many spicules. from New Zealand, A. ciliata Wilson, 1925 from New The proximal end is also frequently sharply bent in the Caledonia, and A. tulearensis Vacelet & Vasseur, 1965 shape of a rhabdostyle. This new species is also notable from Madagascar) or spherical papillate masses (A. for the large overall size of these megascleres. orientalis Dendy, 1905 from Sri Lanka and A. papillata While some authors recognise oxea-like spicules in Lévi & Lévi, 1983 and A. oxytylota Lévi & Lévi, 1983 some species, others appear to lack a second megasclere from New Caledonia). Aciculites spinosa Vacelet & Vas- category. Perhaps a more careful study of all species seur, 1971, also from Madagascar, forms tiny cushions will reveal anatomically differentiated categories. and is found in deep reef tunnels and caves. Key diagnostic charecters: The oxytyles of A. manawatawhi are also quite different from those in other species with their bulbous acan- • Tree-like morphology with small finger-like branch- those heads, but similar to those of A. oxytylota Levi & es of uneven length, surface covered in epizoites. Levi, 1983 from New Caledonia. Like A. manawatawhi, • Deep bluish-gray on the main body, paler branch A. oxytylota also has tubular surface features, but these tips. arise from a cushion base while A. manawatawhi is • Long tylostrongyles with pronounced bulbous columnar with blind digits. heads. Levi and Levi (1983) compared A. oxytylota to the genus Siphonidium (Siphonidiidae), specifically with Aciculites sulcus n. sp. (Fig. 30) S. capitatum Sollas, 1888 from Indonesia, as it also has tubular oscules, but this species is columnar, and more Material examined: similar, in fact, to A. manawatawhi. However, in addition to the similarly shaped ‘oxytylotes’ (Sollas 1888), S. Type material: Holotype NIWA18395 from NIWA Stn capitatum has distinctive planar ectosomal desmas, and KAH9907/48; paratype NIWA18821 from NIWA Stn the oxytyles are exotylote in arrangement — the spicule Z9748. lies perpendicular to the sponge surface with the tylote NIWA Stations: I372 (NIWA25648); RAPUHIA2-15 head facing outwards and the oxeote end pointing into (NIWA25778). the interior (Sollas 1888). Aciculites manawatawhi and A. oxytylota have an irregular but distinct layer of oxytyles Type locality: North Cape and Doubtless Bay; west of that are strewn tangentially and paratangentially in White Island, Bay of Plenty. the surface. This layer is not as thick or distinct as in A. pulchra, for example, but it is definitely present and Distribution: Bay of Plenty. the arrangement is certainly not exotylote. Aciculites manawatawhi and A. oxytylota can also Habitat: Encrusting rocky substrata. Depth range be compared the siphonidiid Gastrophanella, but this 211–250 m. genus also has exotyles [the acanthose subtylostyles

74 Figure 30. Aciculites sulcus n. sp.

75 Description: cific species of Aciculites in that it is cushion-shaped, Morphology of holotype an encrusting arched elongate- with depressed inhalant trough-shaped semicircular rectangular ‘tongue’ attached to the substratum along inhalant areas. In general, the morphology of this new one half of the base (Fig. 30A). Arching along the long- species is quite similar to that of A. orientalis Dendy, est axis of the upper surface of the sponge is a deep 1905, which forms a massive, compact cushion-shaped furrow that contains abundant, regularly dispersed, irregular plate that is attached to calcareous substrata slightly raised, oscules that are c. 0.5 mm diameter. On on a broad base. However, unlike A. sulcus, which one side of the ‘tongue’ is a semicircular indentation presents specialised aquiferous areas, the upper surface that resembles a smooth window that is permeated by of A. orientalis is flakey with oscules and ostia scattered numerous tiny pores or inhalant ostia. The paratype over a slightly corrugated surface (Dendy 1905). Ear- is a fragment of a thick ear-haped semicircular plate shaped specimens are clearly differentiated from A. (Fig. 30B). pulchra Dendy by the absence of raised pimple-like Dimensions 35 mm long, 23 mm at widest point, 15 oscules. mm at the extremities of the sponge, arched 13 mm high Aciculites sulcus is more similar to A. spinosa Vacelet in the centre of the sponge, tapering to about 5 mm at & Vasseur, 1971, a cushion-shaped sponge with spe- each end of the sponge. Furrow is 3–7 mm wide and cialised inhalant areas, but these are circular and the 1–5 mm deep. Inhalant cavity indented c. 5 mm. overall sponge is extremely small (5 mm diameter, 2–3 Texture stony, inner surface of furrow and lateral mm thick), and the tylostrongyles are much smaller indentation flakey. (125–275 µm long, 7–10 µm thick) and acanthose along Surface smooth, granular. the entire spicule. Colour of putty (4B2) in life and in preservative. Key diagnostic characters: Choanosomal skeleton mass of relatively delicate rhizoclone desmas. • Small, thick, arched ear. Ectosomal skeleton of anisostrongyles, arranged ob- • Specialised exhalant furrow along sponge sur- liquely to paratangentially within the upper surface face. furrow and obliquely between the ostia on the surface • Specialised semicircular inhalant depression on side of the lateral inhalant cavity. of sponge. Megascleres comparatively small, delicate rhizoclone • Flush oscules. desmas (Table 27; Fig. 30C,D) with short bifurcate spines and relatively thin clones. Diactinal megascleres Family AZORICIDAE Sollas, 1888 are acanthose anisostrongyles, slightly thinner on the distal end than the proximal end, both ends acanthose, Azoricidae Sollas, 1888: 319; Leiodermatidae Lendenfeld, 1903: 145. proximal end may be occasionally slightly tylote (Fig. 30D). These megascleres are frequently quite sharply Foliose vase-shaped or massive conical sponges with bent in the middle of the spicule and often have a tylote strongly spined rhizoclones; surface diactines may bulge midway along the spicule. emerge beyond sponge surface; microscleres, if present, Microscleres absent. are raphides. (After Pisera and Lévi 2002h). Table 27. Spicule dimensions (µm) of Aciculites sulcus Remarks: Azoricidae Sollas, 1888 is based on very few n. sp. characters and the family contains only two valid Desmas (L) Anisostrongyles (L) genera, recently reviewed by Pisera and Lévi (2002h). Clone Thickness Species of Leiodermatium Schmidt, 1870 have only des- (Thickness) mas and diactinal megascleres, and the single known New Zealand species of Jereicopsis Lévi & Lévi, 1983 has raphide NIWA18395 150–200 228 (210–310) microscleres in addition to desmas and diactines. The (holotype) 20 10–12 type and ornamentation of rhizoclones and the presence of diactinal megascleres suggests a relationship NIWA18821 180–200 300 (210–400) with Microscleroderma in particular (Scleritodermidae) (paratype) 25 15 and this arrangement is supported by 18S rDNA sequence analysis (Kelly-Borges & Pomponi 1994). Pisera Etymology: Named for the specialised exhalant oscu- and Lévi (2002h), however, concluded that, although lar furrow that runs along the surface of the sponge there are similarities, the absence of ectosomal rhabds (Latin, sulcus). and microscleres in Leiodermatium and the presence of Remarks: Aciculites sulcus differs in gross morphology raphides in Jereicopsis distinguish the groups. and spiculation from other Indo- and Southwest Pa-

76 Leiodermatium Schmidt, 1870 Distribution: Patinti Strait between Halmahera and Bacan Islands, Moluccas, Indonesia; Banda Sea; Palau, Leiodermatium Schmidt, 1870: 22; Sollas 1888: 352; Wilson Micronesia. 1925: 465; Pisera & Lévi 2002h: 353. Azorica Carter, 1873: 439. Heterophymia Pomel, 1872: 143. Habitat: Sand and coral boulder slopes. Depth range 27–46 m (Sollas 1888); 375 m (Wilson 1925); 240 m Plate-like, ear-shaped, foliose, or vase-shaped with (Palau). smooth faintly hispid surface; oscules may be visible. Description: Desmas are strongly spined rhizoclones; there are no Morphology a convoluted mass of thin flaring folded special ectosomal spicules and no microscleres. Acces- lamellae that may form tubes, with rounded incised sory megascleres are whip-like diactines. (Modified margins (Wilson 1925; Sollas, 1888) (Fig. 31A,B). The ex- from Pisera and Lévi 2002h). ternal surface of the Palau specimen BMNH 2005.9.15.2 [0CDN 7878-W] has patches of tiny (< 0.5 mm diameter) Type species: Leiodermatium lynceus Schmidt, 1870. ostial pits and the inner surface is completely covered in regularly and closely distributed oscules (< 0.5 mm Remarks: The genus Leiodermatium is best known from diameter). Cup margin in all specimens is thick and the type species Leiodermatium lynceus Schmidt, 1870 rounded, sometimes indented or incised. (Atlantic) and L. pfeifferae (Carter, 1873) and variants Dimensions of Patinti Strait specimens 50–90 mm (Atlantic, Southeast Asia), with additional data be- high, 70–130 mm wide, cup wall 0.5–2.5 mm thick ing provided by much Pacific material (Sollas 1888; (Wilson 1925); Ambon specimen 77 mm high, 110 mm Wilson 1925). wide, lamellae 3.2–4.0 mm thick (Sollas 1888); Palau With relatively few characters available to differen- specimen forms a mass c. 500 mm diameter, wall 2.5 tiate Leiodermatium at the species level, the discovery of mm thick. two new Southwest Pacific taxa — L. linea n. sp. from Texture stony, dense. New Zealand and L. dampieri n. sp. from the Lord Howe Surface smooth and granular with low irregular Seamount Chain — has provided an opportunity to horizontal ridges, internal osculiferous surface slightly examine here the genus as a whole. This revision has re- furry with brishes of projecting diactines. sulted in the recognition of Southeast Asian specimens Colour tan (Palau); topaz (5C5) to putty-like (4B2). of Leiodermatium pfeifferae [Sollas’s (1888) morphotype Skeleton of thorny rhizoclone desmas dominate the ‘Azorica pfeifferae intermedia’ and Leiodermatium pfeifferae choanosome and fine diactinal spicules project from the sensu Wilson (1925)] as L. intermedia n. comb., and in ectosome. Diactines are longer on the margin. the recognition of a new taxon from Micronesia and Megascleres (Table 28) rhizoclone desmas (Fig. the west central pacific, L. colini n. sp. 31C,D) with a straight or curved axis, clones with dis- Leiodermatium intermedia (Sollas, 1888) Table 28. Spicule dimensions (µm) of Leiodermatium (Fig. 31) intermedia (Sollas, 1888).

Azorica pfeifferae intermedia Sollas, 1888: 319, pl. 36, figs Desmas (L x W) Diactines (L) 1–29. Clone Thickness Leiodermatium pfeifferae: Wilson 1925: 465, pl. 43, figs 4, 8, pl. (Thickness) 51, figs 4–7; Lendenfeld 1903: 148. Moluccas, Indonesia Material examined: L. pfeifferae sensu Wilson 1925 Type material: Holotype USNM21321 of Leiodermatium USNM 21321 340 (1) 350–420 pfeifferae (Carter, 1873) and L. pfeifferae f. striata Wilson, (holotype) 35 (2) ? 6 1925; tangential and vertical sections; paratype BMNH 2005.9.15.2, west side of Uchelbeluu Reef, Mutremdiu, Azorica pfeifferae 160–350 (1) ? Koror, Palau, Micronesia (07º16.27’ N, 134º31.37’ E), intermedia Sollas, 1888 20–25 (2) 750 collected by Patrick L. Colin, Coral Reef Research 8 Foundation (Manned submersible Deep Worker 2000), 21 March 2001, 240 m [fragment of 0CDN7878-W]. Palau, Micronesia BMNH 2005.9.15.2 340–500(linear (1) 240–500 forms) Type locality: Ambon Island, Moluccas, Indonesia, 27–46 m (Sollas 1888). (paratype) 35 (2) >1000 1–8

77 Figure 31. Leiodermatium intermedia (Sollas, 1888)

78 crete single or bifurcate spines, the tips of which are — sponges with comparatively thin walls, small closely often bifid, sinuous, or dendritic. On the outer surface spaced oscules, and slender smooth desmas (speci- of the sponge the rhizoclones are smaller and more mens from Bahia); (3) ‘Azorica pfeifferae tenui-laminaris intricate, slightly flattened and with numerous short osculis disjunctis’ — sponges with comparatively thin blunt spines. Diactines (Fig. 31D) in two size categories, walls, small remotely distributed oscules confined to the smallest are fine and hair-like, confined to osculif- the sponge margin (specimen from Bermuda); and erous face of sponge, whereas thicker longer spicules this was differentiated quite clearly from (4) ‘Azorica are confined to the margin of the sponge. pfeifferae intermedia’ — sponges with relatively thick Microscleres absent. walls, small closely spaced oscules, and thick, highly tuberculate desmas (specimen from Ambon Island, Remarks: Leiodermatium pfeifferae Carter, 1873 was first Indonesia). While revision of these Atlantic speci- described from Portugal in the northern Atlantic Ocean, mens is beyond the scope of this work, it appears that from 684 m. Carter (1876) considered L. pfeifferae to be the major differences described are within the range very similar to the type species L. lynceus Schmidt, 1870, of characters expected from sponges collected from also from Portugal, but upheld the separation of his widespread locations. Whether or not the species name new species because of the presence of oscules on the pfeifferae can be applied to these specimens or not will outside of the lamellae rather than on the inside, as in be largely dependent on the ornamentation of the L. pfeifferae and most other planar sponges. Moreover, desmas as other details such as the distribution of the Carter (1876) considered the differences in the distri- diactines are lost. bution of the fine diactines (abundant and restricted Wilson (1925) also attributed several specimens to the margin of L. pfeifferae as a fringe, as opposed to from the 375 m in Patinti Strait, Moluccas, to L. pfeifferae, “isolated diactines around the margin of L. lynceus”) to and these are very similar to the Challenger specimen be significant and thus diagnostic for the two species from nearby Ambon Island. (Table 29). A recent illustration of L. lynceus by Pisera A specimen collected more recently from Palau and Lévi (2002h) shows blunt denticulate spines on the (240 m), by the Coral Reef Research Foundation (Table desmas whereas Sollas (1888) described the desmas 29), is very similar to Wilson’s and Sollas’s Indonesian as terminating in bifurcate spines with bifid rounded material. ends. The ornamentation of these termini is clearly an All of the above Southeast Asian sponges are united important diagnostic indicator of species differences in the possession of a relatively thin-walled foliose (as in other lithistid genera) and careful comparison habit, diactines on all surfaces including the sponge of the ornamentation and morphology of the desmas margin, and large richly tuberculate desmas with using SEM is required to confirm or otherwise the thick clones. They are clearly separate morphologically integrity of L. pfeifferae and L. lynceus. and biogeographically from Atlantic L. pfeifferae, and Sollas (1888) assigned to L. pfeifferae a range of are segregated here as L. intermedia by raising Sollas’s specimens collected by H.M.S. Challenger (1873–1876) (1888) “variety” to full species. Interestingly, this spe- from various Atlantic localities including Bermuda cies is the only known source of two new selectively (795 and 1965 m), Cape Verde Islands (183–234 m), cytotoxic macrocyclic lactones, and these are the first and Brazil (no depth given), but his description of the secondary metabolites reported in the literature to date species (Sollas 1888: 320) was based on a specimen col- (Sandler et al. 2006). lected off Ambon Island in the Moluccas Archipelago, Indonesia (Table 29). This was the only specimen found Key diagnostic characters: to be in the living state upon collection, and Sollas’s • Convoluted mass of thin, flaring, folded lamellae description was based upon this specimen. All of the that may form tubes. Atlantic specimens were dead and denuded skeletons • Oscules and ostia inconspicuous. when collected. • Sponge surface slightly furry. Sollas (1888, p. 322) recognised differences among • Rhizoclones are relatively smooth with numerous the Atlantic specimens and between those and the discrete bifurcate, bifid, dendritic spines. Indonesian specimen. Within the Atlantic complex he recognised at least three morphological ‘groups’ (Table 29) that included (1) ‘Azorica pfeifferae’ — sponges with comparatively thick walls, large marginate oscules, and slender smooth desmas (the holotype specimen of the type species and a specimen from the Cape Verde Islands); (2) ‘Azorica pfeifferae tenui-laminaris’

79 Table 29 (below and opposite). Comparison of Leiodermatium from Atlantic, Southeast Asian, Micronesian, west Central Pacific, and Southwest Pacific waters.

Species Locality and Gross Wall Surface Texture Desma morphology Diactines — depth morphology thickness and size distribution and size

West Central and North Atlantic Leiodermatium lynceus Schmidt, 1870

Type species Portugal and St Ear-shaped 3–4 mm Oscules Stony Smooth with long 190–230 µm, Iago (Carter or hollow large (500– branched spines wispy thin 1876) cylinder 750 µm) and and multidentate (8 µm), with a deep elevated on tips (180–300 µm) isolated on incision in exterior margin margin margin (Schmidt 1870) Leiodermatium pfeifferae (Carter, 1873) Azorica pfeifferae Carter, Madeira, Stalked vase 5 mm Oscules Stony Slender and c. 1785 µm, 1873 Portugal, off with highly small, smooth wispy, thin Holotype Cape St infolded remotely (8 µm), Vincent, 684 m wall distributed restricted to the margin forming a fringe Azorica pfeifferae: Sollas Porta Praya, St Thin plate Compara- Oscules Stony Slender and Unknown 1888 Iago, Cape with tively large and smooth (specimen Verde Islands, infolded thick marginate deciduous) 183–234 m margin (5 mm) Azorica pfeifferae tenui- Bahia, depth Lamella? Compara- Oscules Stony Slender and Unknown laminaris unknown tively thin small and smooth (specimen Sollas, 1888 (3 mm) closely deciduous) distributed Azorica pfeifferae tenui- Bermuda, Lamella? Compara- Oscules Stony Slender and Unknown laminaris osculis disjunctis 795–1965 m tively thin small, smooth (specimen Sollas, 1888 (3 mm) marginate, deciduous) remotely distributed Southeast Asia and Micronesia Leiodermatium intermedia (Sollas, 1888) Azorica pfeifferae intermedia Ambon Island, Thin highly 3.2– Oscules Stony 160–350/20–25 µm 750/8 µm Sollas, 1888 Moluccas, folded plate 4.0 mm small and wide, richly marginate; no Indonesia, with closely tuberculate; spines mention of 27–46 m rounded distributed are bifurcate with smaller margin rounded ends diactines but notes a generally hispid surface. L. pfeifferae sensu Wilson Patinti Strait, Conical, 0.5– Oscules and Stony 340/35 µm, smooth 350-420/3–6 (1925) Moluccas, cup-shaped 2.5 mm ostia clones, termini µm, covering Indonesia, and inconspic- richly tuberculate both surfaces, 375 m lamellate uous with single, but more bifurcate and abundant on bidentate tips concave surfaces BMNH 2005.9.15.2 Palau, 240 m Vase/tube- 2.5 mm Oscules and Stony 340/35 µm, smooth 240–500 µm, forming, ostia clones, termini covering both frilly inconspic- richly tuberculate surfaces, but uous with single, more bifurcate and abundant on bidentate tips concave surface; marginate diactines > 1000 µm 80 Table 29. (continued)

Species Locality and Gross Wall Surface Texture Desma morphology Diactines — depth morphology thickness and size distribution and size

Micronesia and West Central Pacific Leiodermatium colini n. sp.

BMNH 2005.3.14.2 Found in holes Ear- or 8–10 mm Oscules and Stony 100–300/50–100 Very long BMNH 2005.3.14.1 in reef near shallow ostia µm, small desmas diactines Blue Corner, vase-shaped inconspic- with thick clones (> 2000 µm) 28–31 m, West with uous, the and short irregular extending in Barrier Reef, occasional latter in solitary spines tufts from the Palau, undulations meandering margin, Micronesia in lamella; grooves smaller (250– surface and 500 µm) finer margin diactines cover shaggy osculiferous surface only BMNH 2005.3.14.3 Robert’s Reef, 6–8 mm Kimbe Bay, BMNH 2005.3.14.4 West New As above Britain, Papua New Guinea, under overhang, 18–46 m Southwest Pacific and New Zealand Leiodermatium linea n. sp. Bay of Plenty, Frilled cup 3–4 mm Oscules and Stony, Small desmas Fine hair-like New Zealand, (no ostia slightly (120–200)/20 µm; diactines (250– 232–302 m peduncle) inconspic- flexible discrete long single 300 µm) uous, in plane or bifurcate spines aligned in of with bifid or trifid parallel or lamella tips circular channels Leiodermatium dampieri n. sp. Dampier Chalice- to 2.5–4 mm Oscules and Stony 250–350/20–30 µm, 300–450 µm, Ridge, Lord cup- shaped ostia smooth clones, covering both Howe with a short inconspic- termini richly surfaces, but Seamount peduncle uous; ostial tuberculate with more Chain, 420 m pits on some single, bifurcate abundant on parts of the and bidentate tips concave outer surface; surface marginate diactines 675 µm

Leiodermatium colini n. sp. (Fig. 32) June 1997, 31 m [fragment of 0CDN5026-W]; paratype BMNH 2005.3.14.3, Robert’s Reef, Kimbe Bay, West New Britain, Papua New Guinea, 05º25.66’ S, 150º22.65’ Material examined : E, 17 November 1992, 18 m [fragment of 0CDN0779-G]; Type material: Holotype BMNH 2005.3.14.1, cave paratype BMNH 2005.3.14.4, Nomundo Reef, Kimbe off Blue Hole Caverns, West Barrier Reef, Palau, Bay, West New Britain, Papua New Guinea, 05º27.64 S, Micronesia, 07º08.10’ N, 134º13.90’ E, 22 May 2000, 150º06.28’ E, 5 July 2003, 46 m [fragment of 0CDN9062- 28 m [fragment of 0CDN7069-S]; paratype BMNH K]. All specimens collected by P.L. Colin, Coral Reef 2005.3.14.2, cave off Blue Hole Caverns, West Barrier Research Foundation, by SCUBA. Reef, Palau, Micronesia, 07º08.10’ N, 134º13.90’ E, 26

81 Figure 32. Leiodermatium colini n. sp.

82 Distribution: West New Britain (18–46 m), Papua New Microscleres absent. Guinea; Palau (28–31 m), Micronesia. Etymology: Named for Dr Patrick Colin, Director of Habitat: Found on hard bottom in the entrances or the Coral Reef Research Foundation, Palau, Micro- interiors of submarine caves or attached to coral rock nesia, whose deepwater collections and photographs on moderately silty, steep reef slope with overhangs, of lithistid sponges around the Indo-Western Pacific all on fore-reefs experiencing oceanic conditions, 18–46 have significantly increased knowledge of lithistid m; BMNH 2005.3.14.2 and BMNH 2005.3.14.1 were diversity and distribution. found approximately 60 horizontal metres within a cave inside Blue Hole Cavern, Palau, vertical depth Remarks: During the course of this revision, examina- 28–31 m. Depth range 18–46 m. tion of numerous specimens provided by Coral Reef Research Foundation revealed a distinct species group Description: in Palau and from the north coast of Papua New Guinea Morphology of the mature sponge a shallow semicircu- (Table 29). These sponges are comparatively thick- lar plate or cup with broad undulating margins (Fig. walled (c. 10 mm) compared to L. intermedia; they are 32A), or a vertically orientated semicircular plate with ear-shaped or form shallow vases with occasional un- broadly rounded and undulate margin (Fig. 32B). dulations, have 1 mm diameter oscules, and the oscular Dimensions of lamella typically 8–10 mm thick, surface is shaggy with long projecting diactines (> 10 sponge 100–150 mm diameter, BMNH 2005.3.14.3 has mm), particularly on the margin. The desmas are small a thinner lamella (6–7 mm). but the clones are comparatively thick and short, and Texture stony. they are covered in low stubby spines, as opposed to Surface (osculiferous) furry to shaggy in sections the dendritic bifurcate, denticulate spines in L. inter- and on margin, osculiferous face with inconspicuous media. Whilst approaching L. crassiuscula (Sollas, 1888) oscules, < 1 mm diameter, closely regularly scattered from the Kei Islands, Indonesia (256 m) in terms of over entire surface; ostial face with ostia in meandering having a comparatively thick lamella (7 mm), this latter shallow grooves. species forms multiple ear-shaped lamellae, not vases, Colour in life beige (4C3) to tan (6E6), in ethanol does not have a shaggy surface, and the desmas have putty-like (4B2). long multifurcate spines and the desmas are uniaxial. Skeleton of thick heavy rhizoclone desmas dominate The diactines in L. crassiuscula reach 1193 µm. the choanosome (Fig. 21C,D); fine hair-like diactinal The recognition of a second Indo-Pacific species that spicules project from the ectosome on the ostial surface is clearly separate from L. intermedia in terms of gross (Fig. 32C), while thicker diactines form a margin fringe morphology, desma ornamentation, and geographic (Fig. 32D) and dominate the oscular surface. location warrants its inclusion, for comparative pur- Megascleres (Table 30) small rhizoclone desmas with poses, in this monograph of New Zealand taxa. very short thick epirhabd and clones that are covered in low irregular (tuberculate, flattened, incipient bifur- Key diagnostic characters: cation) spines (Fig. 32E,F); diactines with sinuous fine • Thick lamella with a shaggy, fringed, concave sur- termini, in two size categories, the larger confined to face. the margin (Fig. 32D) and the osculiferous surface of • Restricted geographically thus far to Palau and the sponge, the smaller confined to the ostial surfaces northern Papua New Guinea, (Fig. 32F).

Table 30. Spicule dimensions (µm) of Leiodermatium colini n. sp.

Desmas (L x W) Diactines 1 (L) Dactines 2 (L) Clone Thickness Thickness (Thickness) Leiodermatium colini BMNH 2005.3.14.1 150–300 < 5 mm 250–400 (holotype) 50–100 5–13 1–3

BMNH 2005.3.14.2 100–200 5000 300–500 (paratype) 50–80 8–10 1–3

BMNH 2005.3.14.3 150–250 > 1000 300–400 (paratype) 50 8 1–3

83 Leiodermatium dampieri n. sp. (Fig. 33) Table 31. Spicule dimensions (µm) of Leiodermatium dampieri n. sp. Material examined: Desmas (L x W) Diactines (L) Type material: Holotype NIWA18400 from NIWA Clone Thickness Stn P925; paratype NIWA18401, NIWA18402, (Thickness) NIWA18403. Leiodermatium dampieri NIWA Stations: P925 (NIWA18404: 37 specimens from NIWA18400 150–350 (1) 450 the same collection, all dried and mostly deciduous). (holotype) 10–20 (2) 600+

Type locality: Dampier Ridge, Lord Howe Seamount NIWA18402 200–325 none visible Chain. 10–30

Distribution: Dampier Ridge, Lord Howe Seamount NIWA18403 250–425 (1) 300–450 Chain. 20–30 (2) 675 1–8 Habitat: Attached to rocky substrata. Depth range 420 m.

Description: Remarks: The collection of small cup sponges from the Morphology of chalice-shaped or flared plates with a Dampier Ridge in the Lord Howe Seamount Chain is short peduncle by which the sponge attaches to the interesting as it presents a full range of morphologies substratum (Fig. 33A). As the sponge grows in diam- from the smallest chalice-shaped individuals to cups eter, the walls appear to flatten into an undulating plate with flattened undulating walls, yet it is not known and younger specimens are more distinctly chalice- whether these are adults or juveniles. The large number shaped. The base of the peduncle is often flanged and of specimens collected from one location, in one or flared at the point of attachment. The cup margin in all two size classes, suggests that they may be immature specimens is thick and rounded, sometimes indented sponges. or incised. Oscules are inconspicuous but visible under Superficially, these specimens resemble the Asian- magnification and confined to the concave surface, Pacific species L. intermedia and L. colini described excluding the sponge margin (Fig. 33B). in this volume, but careful inspection of the desmas Dimensions — width 10–30 mm wide, overall height reveals that they differ in ornamentation; the desmas 6–20 mm, peduncle height 1–6 mm, peduncle width of L. intermedia are large, arched and thorny with long 8–15 mm; wall thickness 2.5–4 mm. It is not known if bifurcate spines, those of L. colini are quite thick and these are juvenile sponges and grow much larger as coarse with low flat non-bifurcate spines, while those adults. of L. dampieri are more delicate and complex in zygosis, Texture stony, pumice-like (deciduous and dead with rounded protuberances that bifurcate at the very upon collection). tip. The diactines in L. intermedia and L. colini are also Surface smooth and granular, slightly undulating, extremely long compared to those of L. dampieri. convex surface occasionally knobbed or ridged. Until further material of L. dampieri is collected from Colour putty-like (4B2) in the dry state. the remote Dampier Ridge or other seamounts in the Skeleton of thorny rhizoclone desmas that dominate region, it remains to be seen whether these specimens the choanosome; fine diactinal spicules project from include fully mature individuals; the majority are quite the ectosome. small and appear to be juvenile. Megascleres (Table 31) — rhizoclone desmas (Fig. Key diagnostic characters: 33C,D) with a straight or curved axis, clones smooth with occasional rounded to oval prominences, often • Chalice- to plate-shaped tiny sponges. emerging in an angle to the clone, some tubercles bifur- • Rhizoclones are relatively smooth with smooth cate at the extreme tip forming tiny spines. Diactines in rounded protuberances that may bifurcate at the two size categories — the smallest are fine and hair-like, tips. confined to osculiferous face of sponge; thicker longer • May be mistaken for Reidispongia coerulea in the ju- spicules are confined to the margin of the sponge. venile and mature form, but the desma morphology Microscleres absent. is completely different and this species lacks unique megascleres in the surface of the sponge.

Etymology: Named for the type locality.

84 Figure 33. Leiodermatium dampieri n. sp.

85 Leiodermatium linea n. sp. (Fig. 34) Table 32. Spicule dimensions (µm) of Leiodermatium linea n. sp.

Material examined: Desmas (L x W) Diactines (L) Clone Thickness Type material: Holotype NIWA18820 and paratype (Thickness) NIWA18399 from NIWA Stn KAH0011/30; paratype NIWA25048 from TAN0413/118. Leiodermatium linea NIWA18820 170 (120–200) x 250–300 130 (100–150) Type locality: Rungapapa Knoll, Bay of Plenty. (holotype) 20 1

Distribution: Bay of Plenty. NIWA18399 158 (120–200) x 250 100 (80–130 Habitat: Encrusting basalt substrata. Depth range (paratype) 10–20 159–165 m.

Description: Etymology: Named for the aquiferous channels that Morphology (holotype) is a frilled shallow cup with a form visible lines on the lamellae of the sponge, run- slightly expanded rim (Fig. 34A); undulations in the ning from the base to the margin (Latin, linea). wall may anastomose to form a tube. The paratype is a thin basal plate 15 mm diameter and 1 mm thick, from Remarks: The discovery of Leiodermatium in New Zea- which arises a 3 mm thick undulating wall (Fig. 34B). land waters is interesting in that it is the southernmost Oscules and ostia are inconspicuous. Under magnifica- record of the genus, which has not been reported from tion, oscules and ostial pits are c. 0.1 mm diameter. On New Caledonia despite numerous collections on the the concave surface, the oscules are aligned in clearly northern seamounts of the Norfolk Ridge. Although visible shallow aquiferous channels 0.1 mm wide that L. linea is similar in gross morphology to L. intermedia meander from the base of the lamella, stopping short n. sp., subtle differences separate these species (see of the margin. On the convex side, ostial pits are con- Table 29). Leiodermatium linea has characteristic sur- nected in oval interconnecting aquiferous channels. face aquiferous channels that run parallel from the Margin slightly thicker than lamella. base of the lamella to the margin, and are just visible Dimensions 55 mm wide and long, 22 mm high with the unaided eye. On the ostial surface, these lines (holotype); walls 3–4 mm thick. anastomose to form ovals. The desmas and diactines Texture stony. of this species are the smallest yet described, and the Surface smooth, slight growth rings visible, par- ornamentation of the desmas is unique, having short ticularly under the margin, some slight bulges, but conical bifid or trifid spines. otherwise very smooth; slightly furry to the touch with hair-like projecting diactines. Parallel and circular Key diagnostic characters: aquiferous channels visible on concave and convex • Thin-walled cup that may be easily mistaken for surfaces. Reidispongia coerulea, although the latter is blue. Colour in life cream (4A3), putty-like (4B2) in ethanol. Skeleton (choanosome) packed with thorny rhizoclone desmas; hair-like diactinal spicules are arranged radially in the ectosome, protruding slightly from the edge of the sponge. Megascleres (Table 32) — rhizoclone desmas (Fig. 34C) with an arched or straight longitudinal axis; 3–6 clones arise from either side of a straight or curved axis and are covered in well separated discrete single or multiple spines; the apex of these spines may be bifid or trifid but not dendritic (Fig. 34 D,E). Desmas are very fine and comparatively small. Diactines are extremely fine and raphide-like. Microscleres absent.

86 Figure 34. Leiodermatium linea n. sp.

87 DISCUSSION

In total, 282 lithistid specimens were recovered from Caledonian region. Awhiowhio and Lepidothenea are dredge and SCUBA collections resulting from several endemic to New Zealand. hundred NZOI/NIWA voyages in the New Zealand Despite examination of the entire NIWA and EEZ (Gordon 2000). The area covered extends from NZNM collections that span the last 20 years of 24° to 74° S and 155° E to 178° W, covering parts of oceanographic voyages (Gordon 2000), only a single the Lord Howe Rise, Dampier Ridge, South Fiji Basin, lithistid species was recorded south of the Chatham parts of the Norfolk Ridge, the Southwest Pacific Basin, Rise (Neoschrammeniella antarctica from the Ross Sea, Chatham Rise and subantarctic slope, and the Ross Sea, Antarctica), and only a single specimen (Reidispongia Antarctica (Appendix 3). In the north, the area includes coerulea Lévi & Lévi, 1988) was recovered from western Norfolk Island, Lord Howe Island, and the Kermadec New Zealand, on the Challenger Plateau (see Fig. 1). In Islands. Specimens were dredged from 80 to 1680 m, oceanographic terms, New Zealand lithistid sponges but were commonest between 200 and 800 m. Some are not found south of the Subtropical Convergence, of the shallowest sites (140–200 m) were volcanically just south of the Chatham Rise (Shackelton & Kennett active seamounts and knolls in the Bay of Plenty (Run- 1975; Nelson & Cooke 2001), and are thus restricted gapapa, Tuatoru, Tumokemoke, Mahina Knolls) and to relatively ‘warm’ waters on seamounts, continental the crustal rocks of the Cavalli seamount cluster. The margins, and carbonate banks, and to hydrothermally shallowest sites (80 m and 103 m) are off Ngunguru active areas. Bay and in the Cavalli seamounts off Northland. Microfossil spicules very similar to those of living Prior to 1991, only two species of ‘lithistids’ had lithistids have been recorded further south than the been formally reported from New Zealand waters, al- present-day Subtropical Convergence, in the Oamaru though several species were known to exist (Bergquist Diatomite (c. 35 Ma; Hinde & Holmes 1892), and in pers. comm.). The present work recognises 9 families, the Tutuiri Greensand, Chatham Island (c. 56–53 Ma; 18 genera (one of which is new to science), and 30 Kelly & Buckeridge 2005). During the Cenozoic, water species (12 of which are new to science), representing temperatures in the southern parts of the New Zealand a high diversity for this group in a global context. Of region were sufficiently warm to permit establishment the 18 genera, only eight have more than one species of populations of lithistid sponges that today are found (Aciculites, Awhiowhio, Herengeria, Leiodermatium, Neo- only in warmer waters north of the Chatham Rise aulaxinia, Neoschrammeniella, Neosiphonia, and Pleroma), (Campbell et al. 1993). Body-fossils attributed to lithis- confirming the ‘relict’ nature of the group. New spe- tid sponges have also been recorded from the Ototara cies of the previously monospecific phymatellid and Limestone, Kakanui (c. 33.7 Ma; Kelly et al. 2003) and corallistid genera Herengeria, Neoaulaxinia, and Neosi- the Red Bluff Tuff of Chatham Island (Buckeridge & phonia are described, increasing understanding of the Kelly 2006). diagnostic characters of these genera and the extent A preliminary and qualitative comparison of the of their morphological boundaries. Awhiowhio n. gen., spicules of species reported in this work with the closely related to Herengeria in the Corallistidae, was Oamaru Diatomite and Tutuiri Greensand material recognised during the process of character discrimi- of Chatham Island in Kelly et al. (2003), Kelly (2003), nation. This work also records the first polar lithistid and Kelly and Buckeridge (2005) indicates that the species, Neoschrammeniella antarctica, from the Ross Sea. distribution of Recent lithistid sponges differs consid- The description of two new speceis of Leiodermatium in erably from the distribution of fossil lithistids. This New Zealand waters provides the first record of this disjunct distribution between extant and fossil lithistid genus further south than the Philippines. sponges is paralleled in other demosponges and several Fourteen lithistid species are endemic to New Zea- non-siliceous invertebrate groups such as barnacles land waters, 18 are endemic to the New Caledonian (Buckeridge 1983, 1996, 1999), bryozoans (Gordon region, and 14 are ‘regional’ species (Table 33). Several 1984, 1989), and crinoids (Lee 1987; Lee et al. 1997). The genera with species in New Caledonia, viz. Anaderma, typical depth distribution of lithistids today (400–1680 Corallistes, Isabella, Jereicopsis, and Neophrissospongia, m) is in marked contrast to the inferred depth range of are not known in New Zealand waters but are found 50–150 m of lithistid sponges in the warm shallow wa- in Southeast Asia and the western Central Atlantic. ters of southern New Zealand in the Paleogene (35–56 Awhiowhio, Herengeria, and Neoschrammeniella (Coral- Ma; Lee et al. 1997), with the exception of those areas listidae) and Neoaulaxinia, Neosiphonia, and Reidispongia that are today volcanically active, such as the Bay of (Phymatellidae) are endemic to the New Zealand-New Plenty, which harbours almost half of New Zealand’s known lithistid species.

88 Table 33. Checklist of lithistid species described from the New Zealand--New Caledonian region. Note that this list excludes Neoschrammeniella antarctica from the Ross Sea, Antarctica (SEA, Southeast Asia; IO, Indian Ocean).

Species NZ NC Other Aciculites orientalis Dendy, 1905 3 IO Aciculites oxytylota Lévi & Lévi, 1983 3 Aciculites papillata Lévi & Lévi, 1983 3 Aciculites pulchra Dendy, 1924 3 Aciculites sulcus n. sp. 3 Aciculites manawatawhi 3 Anaderma rancureli Lévi & Lévi, 1983 3 Callipelta punctata Lévi & Lévi, 1983 3 3 Corallistes australis Schlacher-Hoenlinger, Pisera & Hooper, 2005 3 Corallistes multituberculatus Lévi & Lévi, 1983 3 Corallistes undulatus Lévi & Lévi, 1983 3 Costifer wilsoni Lévi, 1993 3 3 Discodermia proliferans Lévi & Lévi, 1983 3 3 Awhiowhio osheai n. sp. 3 Awhiowhio sepulchrum n. sp. 3 Awhiowhio unda n. sp. 3 Herengeria auriculata Lévi & Lévi, 1988 3 3 Herengeria vasiformis Schlacher-Hoenlinger, Pisera & Hooper, 2005 3 3 Homophymia pollubrum Schlacher-Hoenlinger, Pisera & Hooper, 2005 3 Homophymia stipitata Kelly, 2000 3 Isabella mirabilis Schlacher-Hoenlinger, Pisera & Hooper, 2005 3 Jereicopsis graphidophora Lévi & Lévi, 1983 3 Leiodermatium dampieri n. sp. 3 Leiodermatium linea n. sp. 3 Lepidothenea incrustans (Dendy, 1924) 3 Macandrewia spinifoliata Lévi & Lévi, 1983 3 3 Microscleroderma herdmani (Dendy, 1905) 3 IO Microscleroderma novaezealandiae n. sp. 3 Microscleroderma stoneae Lévi & Lévi, 1983 3 Neoaulaxinia clavata Lévi & Lévi, 1988 3 3 Neoaulaxinia persicum n. sp. 3 Neoaulaxinia zingiberadix n. sp. 3 Neopelta plinthosellina Lévi & Lévi, 1988 3 Neopelta pulvinus n. sp. 3 Neophrissospongia microstylifer Lévi & Lévi, 1983 3 Neoschrammeniella castrum Schlacher-Hoenlinger, Pisera & Hooper, 2005 3 Neoschrammeniella fulvodesmus Lévi & Lévi, 1983 3 3 Neoschrammeniella moreti (Lévi & Lévi, 1988) 3 Neoschrammeniella norfolkii Schlacher-Hoenlinger, Pisera & Hooper, 2005 3 Neosiphonia superstes Sollas, 1888 3 3 Neosiphonia motukawanui n. sp. 3 3 Pleroma aotea Kelly, 2003 3 Pleroma menoui Lévi & Lévi, 1983 3 3 Pleroma turbinatum Sollas, 1888 3 3 Reidispongia coerulea Lévi & Lévi, 1988 3 3 Scleritoderma camusi Lévi & Lévi, 1983 3 Scleritoderma flabelliformis Sollas, 1888 3 3 SEA TOTAL 29 32 3

89 ACKNOWLEDGMENTS

The author thanks Dr Andrzej Pisera, Institute of Paleo- with information on biological activity of lithistid biology, Polish Academy of Sciences, Warszawa, Po- sponges. land, for his review and comments on the manuscript; Research and fieldwork was funded by a New these have added considerable value to this work. Dr Zealand Lotteries Grants Board Repatriation Fellow- Steve O’Shea, formerly of NIWA Wellington, and Bruce ship (1997–1999), and a Royal Society of New Zealand Marshall, Museum of New Zealand/Te Papa Ton- Marsden Fund Grant (2000–2002). This research has garewa, assisted in sourcing material from the NIWA also been supported by New Zealand Foundation for and NMNZ collections, respectively. The Coral Reef Research Science & Technology Contracts C01X0028 Research Foundation contributed much new material (Seamounts) and C01X0219 (Biodiversity) to NIWA. from their research cruise on NIWA’s RV Kaharoa in This research was also facilitated by the Biological and 1999, around northeastern New Zealand. These and Biotechnological Research Council (BBSRC), UK, and Palau collections were supported by the U.S. National British Airways Conservation Travel Awards, while Cancer Institute shallow-water marine collections pro- the author was employed at the Natural History Mu- gramme. This monograph is thus a contribution of the seum, London (1992–1996). Ongoing support from the Coral Reef Research Foundation. National Institute of Water & Atmospheric Research Dame Professor Patricia Bergquist (retired, Auck- (NIWA) is gratefully acknowledged. land University), provided photographic material for Comparative research on the south New Caledo- several species. Helena Armiger and Lincoln Tubbs nian slope, which assisted this study immensely, was of NIWA Auckland completed all photographic il- carried out onboard the French IRD (Institut de Re- lustrations, and Michael Ellwood of NIWA Hamilton cherche pour le Développement) RV Alis, and I thank prepared the locality map. Drs Alex Ball and Chris Captain Raymond Proner and his crew for their help Jones, Electron Microscopy Unit, Natural History during the voyage. This collaborative work was carried Museum, London were extremely helpful in providing out with financial help of the French Foreign Affair access and training for the author and Klaus Borges, Ministry through the IRD research teams. I particularly UNITEC, Auckland, who carried out the majority of acknowledge the assistance of Dr Bertrand Richer de the scanning microscopy featured in this work. Thanks Forges for his help in the field and Drs Cecile Debitus to Mark Hamann and Jennifer Allman, Pharmacology and Dominique Laurent from the Chemical Product School, University of Mississippi, for their assistance Study Program (SMIB).

REFERENCES

BASSARELLO, C.; NAPOLITANO, A.; BIFULCO, G.; tratigraphy of the Chatham Islands. New Zealand Institute BRUNO, I.; CASAPULLO, A.; RANDAZZO, A.; ZAM- of Geological and Nuclear Sciences Monograph 2: 1–240. PELLA A.; RICCIO, R. 2000: Bioactive macrolides and BUCKERIDGE, J.S. 1996: Phylogeny and biogeography of the cyclopeptides from the South Pacific sponges. Revista primitive Sessilia and a consideration of a Tethyan origin Latinoamericana de Química 28: 59–62. for the group. Crustacean Issues 10: 225–267. BERGQUIST, P.R. 1968: The Marine Fauna of New Zealand: BUCKERIDGE, J.S. 1999: Post Cretaceous biotic recovery: a Porifera, Demospongiae, part 1 (Tetractinomorpha and case study on Crustacea : Cirripedia) from the Chatham Lithistida). New Zealand Oceanographic Institute Memoir Islands, New Zealand. Records of the Canterbury Museum 37: 1–98. 13: 43–51. BEWLEY, C.A.; FAULKNER, D.; ANGEW, J. 1998: Lithistid BUCKERIDGE, J.S.; KELLY, M. 2006: Abstract. On the pal- sponges — star performers or hosts to the stars [re- aeobiology of two distinctive sponge faunas from the view]. Angewanta Chemi International [English Edition] Chatham Islands, New Zealand. 7th International Sponge 37: 2163–2178. Symposium: Biodiversity, Innovation, Sustainability BOURY-ESNAULT, N.; RÜTZLER, K. (Eds) 1997: Thesaurus (7–13 May 2006, Armação de Búzios, Rio de Janiero, of sponge morphology. Smithsonian Contributions to Zool- Brazil). Museu Nacional Série Livros [Published Abstracts] ogy 596: 1–55. 16: 181. BUCKERIDGE, J.S. 1983: The fossil barnacles (Cirripedia: BUCKERIDGE, J.; KELLY, M. 2002: Abstract. Early Cainozoic Thoracica) of New Zealand and Australia. New Zealand hexactinellid sponges (Phylum Porifera) from the Tutuiri Geological Survey Paleontological Bulletin 50: 1–151. Greensand, Chatham Island, New Zealand. Proceedings of the Geological Society of New Zealand Annual Conference, BUCKERIDGE, J.S. 1993: Cirripedia and Porifera. In: Camp- Whangarei, 2–5 December 2002: 10. bell, H.J.; et al. Cretaceous-Cainozoic geology and bios-

90 CAMPBELL, H.J.; ANDREWS, P.B.; BEU, A.G.; MAXWELL, DENDY, A.O. 1924: Porifera. Part I. Non-Antarctic Sponges. P.A.; EDWARDS, A.R.; LAIRD, M.G.; HORNIBROOK, Natural History Report. British Antarctic (“Terra Nova”) N.de B.; MILDENHALL, D.C.; WATTERS, W.A.; BUCK- Expedition, 1910, Zoology 6(3): 269–392, 15 pls. ERIDGE, J.S.; LEE, D.E.; STRONG, C.P.; WILSON, G.J.; DICKEY, J.S. 1968: Observations on the Kakanui Mineral HAYWARD, B.W. 1993: Cretaceous–Cainozoic geology Formation near Kakanui, New Zealand. New Zealand and biostratigraphy of the Chatham Islands. New Zealand Journal of Geology and Geophysics 11: 1159–1162. Institute of Geological and Nuclear Sciences Monograph 2: 1–240. EDWARDS, A.R. 1991: The Oamaru Diatomite. New Zealand Geological Survey Paleontological Bulletin 64: 1–260. CARBONELLI, S.; ZAMPELLA, A.; RANDAZZO, A.; DEBI- TUS, C.; GOMEZ-PALOMA, L. 1999: Sphinxolides E–G GORDON, D.P. 1984: The marine fauna of New Zealand: and reidispongiolide C: four new cytotoxic macrolides Bryozoa: Gymnolaemata from the Kermadec Ridge. New from the new caledonian lithistida sponges N. superstes Zealand Oceanographic Institute Memoir 91: 1–198. and R. coerulea. Tetrahedron 55: 14665–14674. GORDON, D.P. 2000: The Pacific Ocean and global OBIS: a CARTER, H.J. 1876: Descriptions and figures of deep-sea New Zealand perspective. Oceanography 13(3): 41–47. sponges and their spicules from the Atlantic Ocean, GUELLA, G.; MANCINI, I.; DUHET, D.; RICHER de FORG- dredged up on board H.M.S. ‘Porcupine’, chiefly in 1869 ES, B.; PIETRA, F. 1989: Ethyl 6 bromo-3-indolcarboxylate (concluded). Annals and Magazine of Natural History, ser. 4, and 3-hydroxyacetal-6-bromoindole, novel bromoindoles 18: 226–240, 307–324, 388–410, 458–479, pls 12–16. from the sponge Pleroma menoui of the Coral Sea. Zoolo- CHOMBARD, C.; BOURY-ESNAULT, N.; TILLIER, S. 1998: gische Naturforsch, C Bioscience 44: 914–916. Reassessment of homology of morphological characters HARMER, S.F.; LILLIE, D.G.; 1914: List of collecting stations in Tetractinellid sponges based on molecular data. Sys- “Terra Nova” Expedition. Terra Nova Reports, British tematic Biology 47: 351–366. Museum (Natural History) 2: 1–12. CRIST, B.V.; LI, X.; BERGQUIST, P.R.; DJERASSI, C. 1983: HINDE, G.J.; HOLMES, W.M. 1892: On the sponge remains Sterols of marine invertebrates. 44. Isolation, structure in the Lower Tertiary strata near Oamaru, Otago, New elucidation, partial synthesis, and determination of ab- Zealand. Journal of the Linnean Society (London), Zoology solute configuration of pulchrasterol. The first example 24: 177–262. of double bioalkylation of the sterol side chain at position 26. Journal of Organic Chemistry 48: 4472–4479. KELLY, M. 2000a: Description of a new lithistid sponge from northeastern New Zealand, and consideration of DASCH, E.J.; EVANS, A.L.; ESSENE, E. 1970: Radiometric the phylogenetic affinities of families Corallistidae and and petrologic data from eclogites and megacrysts of the Neopeltidae. Zoosystema 22: 265–283. Kakanui Mineral Breccia, New Zealand. Annual Meeting, Geological Society of America, Abstracts with Programs 2 (7): KELLY, M. 2000b: Seamount refuges: sponges show the way. 532–533. Water & Atmosphere 8(3): 7. D’AURIA, M.V.; GOMEZ-PALOMA, L.; MINALE, L.; ZAM- KELLY, M. 2001a: Rock sponges — “living fossils”. Marsden PELLA, A.; DEBITUS, C.; PEREZ, J. 1995: Neosiphoni- Fund Update 16: 5. amolide A, a novel cyclodepsipeptide, with antifungal activity from the marine sponge Neosiphonia superstes. KELLY, M. 2001b: Rock sponges on seamounts: relicts from Journal of natural Products 58: 121–123. the past. Biodiversity Update 4: 3. D’AURIA, M. V.; ZAMPELLA, A.; GOMEZ-PALOMA, L.; KELLY, M. 2003: Revision of the sponge genus Pleroma Sol- MINALE, L.; DEBITUS, C.; ROUSSAKIS, C.; LE BERT, las (‘Lithistida’: Megamorina: Pleromidae) from New V. 1996: Callipeltins B and C; bioactive peptides from a Zealand and New Caledonia, and description of a new marine Lithistida sponge Callipelta sp. Tetrahedron 52: species. New Zealand Journal of Marine and Freshwater 9589–9596. Research 37: 113–127. DAWSON, E.W. 1993: The marine fauna of New Zealand: KELLY, M.; BELL, L.J.; BUCKERIDGE, J.S. 1999: Treasures Index to the fauna: 2. Porifera. New Zealand Oceanographic from the deep: “Living fossil” sponge finds in northern Institute Memoir 100: 1–98. New Zealand are a key to the evolution of the demosponges. Water & Atmosphere 7: 11–12. DENDY, A.O. 1905: Report on the sponges collected by Professor Herdman, at Ceylon, in 1902. In: Herdman, KELLY, M.; BUCKERIDGE, J.S. 2005: An early Paleogene W.A. (ed.), Report to the Government of Ceylon on the Pearl sponge fauna, Chatham Island, New Zealand. New Oyster Fisheries of the Gulf of Manaar 3 (Suppl. 18): 57–246, Zealand Journal of Marine and Freshwater Research 39: 16 pls. 899–914. DENDY, A.O. 1922: Report on the Sigmatotetraxonida col- KELLY, M.; EDWARDS, A.R.; WILKINSON, M.R.; ALVAR- lected by the H.M.S. ‘Sealark’ in the Indian Ocean. In: EZ, B.; COOK, S.deC.; BERGQUIST, P.R.; BUCKERIDGE, Reports of the Percy Sladen Trust Expedition to the Indian J.S.; CAMPBELL, H.; REISWIG, H.M.; VALENTINE, C. In Ocean in 1905, Volume 7. Transactions of the Linnean Society press: Phylum Porifera—sponges. In: Gordon, D.P. (ed.), of London, ser. 2, 18: 1–164, 18 pls. The New Zealand Inventory of Biodiversity Volume 1. King- dom Animalia: Radiata, Lophotrochozoa, and Deuterostomia. Canterbury University Press, Christchurch.

91 KELLY, M. 2004: News: The oceans’ own ‘tuatara’ discovered. Crosnier, A. (ed.), Résultats des campagnes Musorstom Water & Atmosphere 11(4): 5. 11. Mémoires du Muséum nationale d’Histoire naturelle, sér. A, Zoologie, 158: 9–87. KELLY, M.; LEE, D.; KELLY, S.: BUCKERIDGE, J.S. 2003: A Recent sponge, Pleroma aotea Kelly (‘order’ Lithistida: LÉVI, C.; LÉVI, P. 1983: Éponges tetractinellides et lithistides family Pleromidae), in the Late Eocene Ototara Limestone bathyales de Nouvelle Calédonie. Bulletin du Muséum of Otago, New Zealand. New Zealand Journal of Marine and national d’Histoire naturelle, sér. 4, 5(A,1): 101–168. Freshwater Research 37: 129–148. LÉVI, C.; LÉVI, P. 1988: Nouveaux Spongiaires — éponges KELLY, M.; TUBBS, L.2006: The lithistid Demospongiae of lithistides bathyaux à affinitiés crétacées de la Nouvelle- New Zealand. 7th International Sponge Symposium: Calédonie. Bulletin du Muséum national d’Histoire naturelle, Biodiversity, Innovation, Sustainability (7–13 May 2006, sér. 4, 10(A,2): 241–263. Armação de Búzios, Rio de Janiero, Brazil). Museu Na- LÉVI, C.; LÉVI, P. 1989: Spongiaires (Musorstom 1 & 2). In: cional Série Livros 16 [Published abstracts]: 288. Forest, J. (ed.) Résultats des Campagnes Musorstom, 4. KELLY-BORGES, M. ROBINSON, E.V.; GUNASEKERA, S.; Mémoires du Muséum nationale d’Histoire naturelle, sér. A, GUNASEKERA, M.; GULAVITA, N.; POMPONI, S.A. 143: 25–103 1994: Species differentiation in the marine sponge genus MAYER, A.M.S.; HAMANN, M.T. 2004: Marine pharmacol- Discodermia (Demospongiae, Lithistida): The utility of ogy in 2000: marine compounds with antibacterial, anti- secondary metabolites as species-specific markers. Bio- coagulant, antifungal, anti-inflammatory, antimalarial, chemical Systematics and Ecology 22: 353–365. antiplatelet, antituberculosis, and antiviral activities; KELLY-BORGES, M.; POMPONI, S.A. 1994: Phylogeny and affecting the cardiovascular, immune and nervous Re- classification of lithistid sponges (Porifera: Demospong- systems and other miscellaneous mechanisims of action. iae): a preliminary assessment using ribosomal DNA Marine Biotechnology 6: 37–52. sequence comparisons. Molecular Marine Biology and McINERNEY, J.O.; ADAMS, C.L.; KELLY, M. 1999: Phylo- Biotechnology 3: 87–103. genetic resolution potential of 18S and 28S rRNA genes KELLY-BORGES, M.; VACELET, J. 1995: A revision of Diacar- within the lithistid Astrophorida. Memoirs of the Queens- nus Burton and Negombata de Laubenfels (Demospongiae: land Museum 44: 343–352. Latrunculiidae) with descriptions of new species from MORET, L. 1926: Contribution à l’étude des Spongiaires the West Central Pacific and the Red Sea. Memoirs of the silicieux de Crétacé supérieur français. Mémoires de la Queensland Museum 38: 477–503. Société Géologique de France 5: 1–267. KORNERUP, A.; WANSCHER, J.H. 1961: Reinhold Color Atlas. MUNRO, M.H.G.; BLUNT, J.W.; DUMDEI, E.J.; HICKFORD, Reinhold Publishing Corporation, New York. 224 p. S.J.H.; LILL, R.E.; LI, S.; BATTERSHILL, C.N.; DUCK- LAILLE, M.; GERALD, F.; DEBITUS, C. 1998: In vitro antiviral WORTH, A.R. 1999: The discovery and development activity on dengue virus of marine natural products. Cel- of marine compounds with pharmaceutical potential. lular and Molecular Life Sciences 54: 167–170. Journal of Biotechnology 70: 15–25.

LAUBENFELS, M.W. de 1936: A discussion of the sponge NELSON, C.S.; COOKE, P.J. 2001: History of oceanic front fauna of the Dry Tortugas in particular and the West development in the New Zealand sector of the Southern Indies in general, with material for the revision of the Ocean during the Cenozoic — a synthesis. New Zealand families and orders of the Porifera. Publications of the Journal of Geology and Geophysics 44: 535–553. Carnegie Institution, No. 467, Papers from the Tortugas PIEL, J.; HUI, D.; WEN, G.; BUTZKE, D.; PLATZER, M.; Laboratory 30: 1–224. FUSETANI, N.; MATSUNAGA, S. 2004: Antitumor LAUBENFELS, M.W. de 1955: Porifera. Pp. 21–112 in: Moore, polyketide biosynthesis by an uncultivated bacterial sym- R.C. (ed.), Treatise on Invertebrate Paleontology, Part E. Ar- biont of the marine sponge Theonella swinhoei. Proceedings chaeocyatha and Porifera, Geological Society of America and of the National Academy of Sciences 101: 16222-16227. University of Kansas Press, Boulder & Lawrence. PISERA, A. 1999: Lithistid sponge Setidium obtectum Schmidt, LEE, D. 1987: Cenozoic and Recent inarticulate brachiopods 1879, rediscovered. Memoirs of the Queensland Museum of New Zealand: Discinisca, Pelagodiscus and Neocrania. 44: 473–477. Journal of the Royal Society of New Zealand 17: 49–72. PISERA, A. 2002: Fossil lithistids: an overview. Pp. 388–402 in: LEE, D.E.; SCHOLZ, J.; GORDON, D.P. 1997: Paleoecology Hooper, J.N.A.; Soest, R.W.M. van (eds), Systema Porifera: of a Late Eocene mobile rockground biota from north Guide to the Classification of Sponges. Kluwer Academic/ Otago. Palaios 5: 568–581. Plenum Publishers, New York.

LEHNERT, H.; SOEST, R.W.M. van 1996: North Jamaican PISERA, A. 2006: Palaeontology of sponges — a review. deep fore-reef sponges. Beaufortia 46: 53–81. Canadian Journal of Zoology 84: 242–261. LÉVI, C. 1991: Lithistid sponges from the Norfolk Rise: Recent PISERA, A.; LÉVI, C. 2002a: lithistid Demospongiae. Pp. and Mesozoic genera. Pp. 72–82 in: Reitner, J.; Keupp, H. 299–301 in: Hooper, J.N.A.; Soest, R.W.M. van (eds), Sys- (eds), Fossil and Recent Sponges. Springer-Verlag, Berlin. tema Porifera: Guide to the Classification of Sponges. Kluwer Academic/Plenum Publishers, New York. LÉVI, C. 1993: Porifera Demospongiae: Spongiaires bathyaux de Nouvelle-Calédonie, récoltés par le ‘Jean Charcot’. In: 92 PISERA, A.; LÉVI, C. 2002b: Family Scleritodermidae Sollas, RICHER de FORGES, B.; KOSLOW, J.-A.; POORE, G.C.B. 1888. Ibid: 302–311. 2000: Diversity and endemism of the benthic seamount macrofauna in the southwest Pacific. Nature 405: PISERA, A.; LÉVI, C. 2002c: Family Corallistidae Sollas, 1888. 944–947. Ibid: 312–320. SANDLER, J.S.; COLIN, P.L.; KELLY, M.; FENICAL, W. 2006: PISERA, A.; LÉVI, C. 2002d: Family Pleromidae Sollas, 1888. Cytotoxic macrolides from a new species of deep-water Ibid: 321–326. marine sponge Leidermatium. Journal of Organic Chemistry PISERA, A.; LÉVI, C. 2002e: Family Theonellidae Lendenfeld, 71: 7245–7251. 1888. Ibid: 327–337. SCHLACHER-HOENLINGER, M.A.; PISERA, A.; HOOPER, PISERA, A.; LÉVI, C. 2002f: Family Siphonidiidae Lenden- J.N.A. 2005: Deep-sea “lithistid” assemblages from the feld, 1888. Ibid: 338–343. Norfolk Ridge (New Caledonia), with description of seven new species and a new genus (Porifera, Demos- PISERA, A.; LÉVI, C. 2002g: Family Neopeltidae Sollas, 1888. pongiae). Zoosystema 27: 649–698. Ibid: 344–351. SCHMIDT, O. 1870: Gundzüge einer Spongien Fauna des atlan- PISERA, A.; LÉVI, C. 2002h: Family Azoricidae Sollas, 1888. tischen Gebietes. Wilhelm Engelman, Leipzig. 88 p. Ibid: 352–355. SCHMIDT, O. 1880: Die Spongien des Meerbusens von Mexico PISERA, A.; LÉVI, C. 2002i: Family Desmanthidae Topsent, und des Caraibischen Meeres. Gustav Fischer, Jena. 90 p. 1888. Ibid: 356–362. SCHRAMMEN, A. 1910: Die Kieselspongien der oberen PISERA, A.; LÉVI, C. 2002j: Family Vetulinidae Lendenfeld, Kreide von Nordwestdeutschland. Tetraxonia, Monaxo- 1888. Ibid: 363–365. nia und Silicea. Grundzüge einer Spongien fauna des At- PISERA, A.; LÉVI, C. 2002k: Family Phymatellidae Schram- lantischen Gebietes. Palaeontographica, Suppl. 5(1): 1–84. men, 1888. Ibid: 366–373. SHACKLETON, N.; KENNETT, J. 1975: Paleotemperature PISERA, A.; LÉVI, C. 2002l: Family Isoraphiniidae Schram- history of the Cenozoic and the initiation of Antarctic men, 1888. Ibid: 374–376. glaciation: oxygen and carbon isotope analyses in DSDP Sites 277, 279 and 281. In: Kennett, J.; Houtz, R. (eds). PISERA, A.; LÉVI, C. 2002m: Family Macandrewiidae Initial Reports of the Deep Sea Drilling Program, Washington Schrammen, 1888. Ibid: 377–379. 29: 743–755. PISERA, A.; LÉVI, C. 2002n: Family Phymaraphinidae SOEST, R.W.M. van; STENTOFT, N. 1988: Barbados deep- Schrammen, 1888. Ibid: 380–383. water sponges. Studies on the Fauna of Curaçao and other PISERA, A.; LÉVI, C. 2002o: Lithistids incertae sedis. Ibid: Caribbean Islands 70: 1–175. 384–387. SOLLAS, W.J. 1888: Report on the Tetractinellida collected by POMPONI, S.A. 2001: The oceans and human health: the H.M.S. Challenger during the years 1873–1876. Reports on discovery and development of marine-derived drugs. the Scientific Results of the Voyage of the H.M.S. Challenger, Oceanography 14: 78–87. Zoology 25(63): 1–458. POMPONI, S.A.; KELLY, M.; REED, J.; WRIGHT, A. 2001: Di- SUGGATE, R.P.; STEVENS, G.R.; TE PUNGA, M.T. (Eds) versity and bathymetric distribution of lithistid sponges 1978: The Geology of New Zealand.Government Printer, in the tropical western Atlantic region. Bulletin of the Wellington. 2 vols, 820 p. Biological Society of Washington 10: 344–353. VACELET J.; VASSEUR, P. 1965: Spongiaires des grottes et REED, J.K.; POMPONI, S.A. 1997: Biodiversity and distribu- surplombs des récifs de Tuléar (Madagascar). Recueil des tion of deep and shallow water sponges in the Bahamas. travaux de la Station marine d’Endoume 4: 71–123. Proceedings of the Eighth International Coral Reef Symposium VACELET, J., VASSEUR, P. 1971: Éponges des Récifs coral- 2: 1387–1392. liens de Tuléar (Madagascar). Tethys, Suppl. 1: 56–126. REID, R.E.H. 1963: A classification of the Demospongia. WILSON, H.V. 1925. Siliceous and horny sponges collected Neues Jahrbuch für Geologie und Paläontologie. Monatshefte by the U.S. Fisheries Steamer ‘Albatross’ during the 4: 196–207. Philippine Expedition, 1907–1910. In: Contributions to REID, R.E.H. 1970: Tetraxons and demosponge phylogeny. the biology of the Philippine Archipelago and adjacent Pp. 63–89 in: Fry, W.G. (ed.), The Biology of the Porifera. regions. Bulletin of the United States National Museum 100: [Symposia of the Zoological Society of London 25.] Aca- 273–532. demic Press, London. ZAMPELLA, A.; D’AURIA M.V.; MINALE, L.; DEBITUS, REID, R.E.H. 2004: Mesozoic and Cenozoic lithisitd sponges: C. 1997: Callipeltosides B and C, two novel cytotoxic Dicranocladina, Pseudorhizomorina, Didymorina, glycoside macrolides from a marine lithistida sponge Helomorina, Megamorina, Megarhizomorina, Sphaero- Callipelta sp. Tetrahedron 53: 3243–3248. cladina and order and suborder uncertain. Pp. 239–274 ZITTEL, K.A. von 1878: Studien über fossile Spongien. Zweite in: Kaessler, R.E. (ed.), Treatise on Invertebrate Palaeontol- Abtheilung: Lithistidae. Abhandlungen der K. Bayer Akad- ogy. Part E (revised). Porifera. Vol 3. Geological Society of emie der Wissenschaft 13: 65–154. America & University of Kansas, Boulder & Lawrence.

93 Appendix 1

Key to diagnostic characters for massive, spherical, club-shaped and small encrusting lithistid sponges in New Zealand waters This key differentiates massive, spherical, and club-shaped sponges, or small encrusting or cushion-shaped sponges on their gross morphology, a character that can be used reasonably reliably in the field. Where the morphology of species is similar, diagnostic microscopic characters (mic) are given for a final confirmation. For very closely related species in the genera Neoaulaxinia and Neosiphonia, a full microscopic examination of the spicules will be required.

Small stony nubbin sponges with tufts of long spicules emerging from the top (Fig. 11A,B) Herengeria auriculata Small nodulose branches or irregular tuberose mass...... 1 1. Stony texture with surface discotriaenes (mic) (Fig. 2A,D) ...... Discodermia proliferans Comparatively soft with short-rhabd dichotriaenes (mic) (Fig. 4A,B)...... Neoaulaxinia zingiberadix Club-shaped with apical region expanding from a narrow base...... 2 2. Parsnip-shaped with a grey-blue apical oscular depression (Fig. 16A,B)...... Homophymia stipitata Thin elongate club with a single oscule at the apex (Fig. 3A,B)...... Neoaulaxinia clavata Tree-like with small finger-like branches of uneven length, surface covered in epizoites (Fig. 29A–C)...... Aciculites manawatawhi Small mushroom-shaped bulbous body with an apical depression and short stalk...... 3 3. Small stony squat spherical body with a restricted base and slightly concave bumpy surface (Fig. 21D,E)...... Pleroma aotea Squat body elevated on a short stem, deep roofed apical depression, comparatively soft and crumbly (Fig. 6A)...... Neosiphonia superstes Egg-shaped, to spherical block-like masses, surface flakey, granular or conulose in places...... 4 4. Spherical to egg-shaped with a single apical oscule, short dichotriaene rhabd (mic) (Fig. 5A–C)...... Neoaulaxinia persicum Typically block-shaped with apical cavity and ‘parchment’ roof, long dichotriaene rhabd (mic.) (Fig. 7A,B) ...... Neosiphonia motukawanui Tiny nodules and cushions...... 5 5. Columnar sponges (<20 mm diamater) with apical oscule, with surface pseudophyllotriaenes (mic) (Fig. 17A)...... Callipelta punctata Cushion-like encrustation with surface scale-like discs (mic) (Fig. 18A)...... Neopelta pulvinus Thin flat silvery encrustation...... 6 6. Surface has feather-like triaenes and curved strongylote oxeas (mic) (Fig. 20 C,D)...... Macandrewia spinifoliata Surface has scale-like discs and acanthose microrhabds (mic) (Fig. 19)...... Lepidothenea incrustans

94 Appendix 2

Key to diagnostic characters for cup-, vase-, tube- or plate-shaped lithistid sponges in New Zealand waters

This key differentiates cup-, vase-, funnel-, tube-, plate- (flat or frilled), and ear-shaped sponges on key diagnostic spicule characteristics rather than on gross morphology, because the shape of these sponges is not a reliable field character. In most cases, immature sponges of all species have very similar shapes and many cup-shaped sponges occur in an ear-shaped or plate form and vice versa. While some species have characteristic surface features (pimpled, dimpled, lines of oscules, concentric lines across the face, and thick or thin margin) and textures (velvety, granular, fringe of spicules), these characters are also generally unreliable, unless in combination with particular spicule characteristics. The presence (or absence) of diagnostic spicule types differentiates between genera in this key, and the range of shapes and sizes within the microsclere categories differentiates species within a genus. Where a particular morphological or surface characteristic is diagnostic in combination with the primary spicule characters (e.g. in Reidispongia coerulea), this character is also employed.

Microscleres include two types of amphiasters (Fig. 8F), mature sponge a vivid blue frilly mass (Fig. 8A,B)...... Reidispongia coerulea Microscleres include sigmaspires...... 1 1. Surface crust of acanthorhabds present (Fig. 26B)...... Scleritoderma flabelliformis Acanthorhabds absent (Fig. 25C)...... Microscleroderma novaezelandiae Megascleres include tylostrongyles with roughened ends...... 2 2. Tongue- or ear-shaped, oscules flush with aquiferous ‘slits’ emanating from oscule (Fig. 30).. Aciculites sulcus Oscules raised on tubercles on one surface, reddish-brown (Fig. 27A-D)...... Aciculites pulchra Thick, furry ear (Fig. 23 A); desmas are long, notched heloclones (Fig. 24)...... Costifer wilsoni Desmas are smooth megaclones with saddle-shaped zygomes...... 3 3. Desmas are distinctive smooth ‘fat’ dipods, tripods, or quadripods (Fig. 22B,C,F)...... Pleroma menoui Desmas are slim smooth ‘slender’ quadripods or multipods (Fig. 22A,E), usually a thick stony ear sponge (Fig. 21A,B)...... Pleroma turbinatum Microscleres include two types of spiraster (robust and fine-rayed) and a very large fine-rayed streptaster...... 4 4. Thin-walled (4–8 mm) shallow or tall cup-shaped sponges with flared margin (Fig. 9A–C)...... Neoschrammeniella fulvodesmus Thick-walled tall conical cup with fringed internal margin (Ross Sea only) (Fig. 10A)...... Neoscrammeniella antarctica Microscleres include 2 types of spiraster (robust and fine-rayed), an ectosomal polyrhabd and acanthose centrotylote microxeas...... 5 5. Polyrhads very irregular (Fig. 12G)...... Herengeria vasiformis Polyrhabds centrotylote with bulbous ends (Fig. 13G), acanthose microxeas absent, one surface pimpled (Fig. 13A–C)...... Awhiowhio osheai Polyrhabds spiraster-shaped, acanthose microxeas absent, sponge with wavy walls (Fig. 14A,D,E)...... Awhiowhio unda Sponge a large thick flat plate (Fig. 15)...... Awhiowhio sepulchrum Microscleres absent...... 6 6. Thin-walled cup with parallel vertical lines on the surface (Fig. 34A)...... Leiodermatium linea

95 Appendix 3 Station data

NIWA/NZOI (New Zealand Oceanographic Institute) Stations

NIWA Stn Geographic location Latitude Longitude Depth (m) Date A527 Ross Sea, Antarctica 74°10.00’ S 178°17.0’ W 352 02.07.1960 B314 Eastern edge of Challenger Plateau 39°22’ S 171°50.0’ E 236 25.10.1960 E636 Bay of Plenty 37°28.49’ S 177°13.0’ E 190 10.10.1966 E731 Bay of Plenty 37°23.50’ S 177°12.0’ E 602–503 25.03.1967 F993 South Three Kings Ridge 34°24’ S 173°10.3’ E 249–212 15.10.1968 I14 Cavalli Seamount, Northland 35°35.9’ S 174°39.7’ E 103 04.05.1975 I92 Norfolk Ridge, south of Norfolk Island 29°24.8’ S 168°13.2’ E 578–570 23.07.1975 I96 Wanganella Bank, West Norfolk Ridge 32°10.8’ S 167°21.2’ E 356 05.02.1975 I97 Wanganella Bank, West Norfolk Ridge 32°22.9’ S 167°28.2’ E 544–540 25.07.1975 I372 Doubtless Bay, North Cape 34°32.20’ S 173°29.50’ E 215–211 23.11.1977 J953 (I807) Western continental slope, Northland 34°39.6’ S 172°13.1’ E 270–260 18.06.1981 J954 (I808) Western continental slope, Northland 34°38.8’ S 172°13.5’ E 204–192 18.06.1981 K826 Southern South Fiji Basin 28°48.0’ S 177°48.0’ W 490–390 25.07.1974 K872 Colville Ridge, southeastern South Fiji Basin 31°20.4’ S 178°49.2’ W 280–235 02.08.1974 KAH0011/30 Rungapapa Knoll, Bay of Plenty 37º32.98’ S 176°58.84’ E 165–159 04.11.2000 KAH0011/40 Rungapapa Knoll, Bay of Plenty 37°32.99’ S 176°58.25’ E 280–155 05.11.2000 KAH0011/41 Rungapapa Knoll, Bay of Plenty 37°33’ S 176°58.23’ E 260–154 05.11.2000 KAH0011/44 Rungapapa Knoll, Bay of Plenty 37°33.14’ S 176°58.09’ E 348–181 05.11.2000 KAH0204/02 Knights Terrace, east of Poor Knights Islands 34°57.62’ S 175°10.79’ E 581 13.04.2002 KAH0204/07 Cavalli Seamount, Northland 34°7.16’ S 174°9.15’ E 800 14.04.2002 KAH0204/08 Cavalli Seamount, Northland 34°6.91’ S 174°8.70’ E 640–610 14.04.2002 KAH0204/09 Cavalli Seamount, Northland 34°6.68’ S 174°8.29’ E 600–562 14.04.2002 KAH0204/27 Cavalli Seamount, Northland 33°09.39’ S 179°57.2’ E 667–613 16.04.2002 KAH0204/29 West Cavalli Seamount, Northland 34°9.79’ S 173°57.75’ E 790–782 17.04.2002 KAH0204/30 West Cavalli Seamount, Northland 34°8.79’ S 173°57.79’ E 825–800 17.04.2002 KAH0204/32 West Cavalli Seamount, Northland 34°9.72’ S 173°57.71’ E 810–780 17.04.2002 KAH0204/38 West Cavalli Seamount, Northland 34°09.52’ S 173°57.78’ E 800–780 18.04.2002 KAH0204/40 West Cavalli Seamount, Northland 34°09.85’ S 173°57.84’ E 820–805 18.04.2002 KAH0204/44 South Cavalli Seamount, Northland 34°15.94’ S 174°06.19’ E 850–840 18.04.2002 KAH0204/47 Seamount 441, Cavalli Seamount region 34°02.55’ S 174°49.02’ E 880–792 19.04.2002 KAH0204/52 Seamount 441, Cavalli Seamount region 34°03.30’ S 174°48.49’ E 910–820 19.04.2002 KAH9301 Bay of Plenty to Wairarapa shelf 36°57.24’ S 176°15.19’ E 31.12.1992 to 41°01.62’ S 176°24.87’ E 27.01.1993 KAH9907/48 West of White Island, Bay of Plenty 37°28.15’ S 177°06.71’ E 250 05.06.1999 KAH9907/49 West of White Island, Bay of Plenty 37°28.21’ S 177°07.03’ E 320–220 05.06.1999 KAH9907/50 West of White Island, Bay of Plenty 37°28.14’ S 177°06.97’ E 318–230 05.06.1999 P8 West Norfolk Ridge 32°40.8’ S 167°26.80’ E 757–660 25.01.1977 P10 West Norfolk Ridge 32°40.00’ S 167°28.40’ E 378–352 25.01.1977 P14 West Norfolk Ridge 31°47.20’ S 167°51.6’ E 319–316 25.01.1977 P925 Dampier Ridge, Lord Howe Seamount Chain 27°59.60’ S 155°37.50’ E 420 11.12.1979 RAPUHIA2-15 [NIWA Stns U545–U624, cruise to study manganese crusts; depths 406–3354 m] Challenger Plateau to southern Norfolk Ridge 40°16.5’ S 165°59.0’ E 29.01.1988 to 30°20.0’ S 172°60.0’ E 14.02.1988 S562 Western continental slope, Northland 35°49.20’ S 172°54.00’ E 600–505 05.08.1983 S568 Western flank of Three Kings Ridge 30°10.00’ S 171°20.20’ E 900–650 13.08.1983 S571 Three Kings Ridge 30°47.30’ S 172°45.20’ E 509–480 15.08.1983 S572 West of Three Kings Ridge 30°45.50’ S 172°47.70’ E 530–403 15.08.1983 T226 North of Raoul Island, Kermadec Ridge 28°33.00’ S 177°49.99’ W 800–930 22.03.1982 TAN0104/001 Graveyard Seamount, Chatham Rise 42°45.60’ S 179°59.26’ W 979–770 15.04.2001 TAN0104/002 Graveyard Seamount, Chatham Rise 42°45.93’ S 179°59.34’ W 875–757 15.04.2001 TAN0104/048 Diabolical Seamount, Chatham Rise 42°47.17’ S 179°59.12’ W 993–900 16.04.2001 TAN0104/148 Morgue Seamount, Chatham Rise 42°42.84’ S 179°57.51’ W 980–893 18.04.2001 TAN0104/194 Scroll Seamount, Chatham Rise 42°47.27’ S 179°59.81’ W 1042–880 18.04.2001 TAN0104/288 Graveyard Seamount, Chatham Rise 42°45.64’ S 179°59.27’ W 972–890 19.04.2001

96 NIWA/NZOI (New Zealand Oceanographic Institute) Stations

NIWA Stn Geographic location Latitude Longitude Depth (m) Date TAN0104/289 Graveyard Seamount, Chatham Rise 42°45.89’ S 179°59.16’ W 800–757 19.04.2001 TAN0104/394 Graveyard Seamount, Chatham Rise 42°45.68’ S 179°59.33’ W 920–771 21.04.2001 TAN0107/124 Rumble III Seamount, Bay of Plenty 35°44.36’ S 178°30.25’ E 620–435 20.05.2001 TAN0107/232 Rumble V Seamount, Bay of Plenty 36°08.73’ S 178°11.98’ E 750–570 24.05.2001 TAN0205/20 Volcano E, Kermadec Ridge 33°44.18’ S 179°49.90’ W 619–490 14.04.2002 TAN0205/32 East slope of Kermadec Ridge 33°10.25’ S 179°58.21’ W 999–643 16.04.2002 TAN0205/60 Cavalli Seamount, Northland 30°11.87’ S 178°28.75’ W 1131–1073 21.04.2002 TAN0205/75 Volcano L, Kermadec Ridge 30°01.37’ S 178°42.88’ W 240–154 23.04.2002 TAN0413/016 Whakatane Seamount, Bay of Plenty 36°47.50’ S 177°25.52’ E 1538–1503 08.11.2004 TAN0413/068 Tuatoru Knoll, Bay of Plenty 37°28.28’ S 177°12.30’ E 295–200 11.11.2004 TAN0413/070 Tuatoru Knoll, Bay of Plenty 37°28.12’ S 177°12.40’ E 330–218 11.11.2004 TAN0413/074 Tuatoru Knoll, Bay of Plenty 37°28.19’ S 177°13.20’ E 200–175 12.11.2004 TAN0413/077 Tuatoru Knoll, Bay of Plenty 37°28.26’ S 177°12.89’ E 180–177 12.11.2004 TAN0413/085 Tuatoru Knoll, Bay of Plenty 37°28.25’ S 177°13.35’ E 216–207 12.11.2004 TAN0413/092 Tuatoru Knoll, Bay of Plenty 37°29.14’ S 177°12.92’ E 400–225 12.11.2004 TAN0413/093 Tuatoru Knoll, Bay of Plenty 37°28.58’ S 177°12.94’ E 180–179 13.11.2004 TAN0413/099 Rungapapa Knoll, Bay of Plenty 37°32.80’ S 176°58.85’ E 151–141 13.11.2004 TAN0413/109 Rungapapa Knoll, Bay of Plenty 37°32.93’ S 176°59.27’ E 142–136 13.11.2004 TAN0413/111 Rungapapa Knoll, Bay of Plenty 37°32.79’ S 177°00.22’ E 337–256 13.11.2004 TAN0413/112 Rungapapa Knoll, Bay of Plenty 37°32.11’ S 176°58.26’ E 217–169 13.11.2004 TAN0413/117 Rungapapa Knoll, Bay of Plenty 37°32.16’ S 176°58.25’ E 188–182 13.11.2004 TAN0413/118 Rungapapa Knoll, Bay of Plenty 37°33.22’ S 176°58.13’ E 190–154 13.11.2004 TAN0413/120 Mahina Knoll, Bay of Plenty 37°21.14’ S 177°04.70’ E 520 14.11.2004 TAN0413/170 Tumokemoke Knoll, Bay of Plenty 37°27.69’ S 176°54.23’ E 376–295 16.11.2004 TAN0413/173 Mahina Knoll, Bay of Plenty 37°21.04’ S 177°05.96’ E 434–272 16.11.2004 TAN0413/174 Mahina Knoll, Bay of Plenty 37°20.09’ S 177°04.58’ E 502–430 16.11.2004 TAN9915/3B Ngunguru Bay, off Taiharuru Head, Northland 35°41.80’ S 174°38.90’ E 80 13.12.1999 U591 Tui Seamount, Three Kings Ridge 30°50.60’ S 172°48.30’ E 486 07.02.1988 U594 West of Three Kings Ridge 30°20.10’ S 172°59.60’ E 406 07.02.1988 U606 South Norfolk Basin 31°54.70’ S 172°47.40’ E 1100–1085 10.02.1988 V473 South of East Cape 38°50.92’ S 178°36.75’ E 800–365 05.06.1994 X128 Bay of Plenty 37°24.40’ S 176°59.40’ E 405 25.11.1989 X140 Bay of Plenty 37°11.40’ S 176°58.30’ E 860–745 27.11.1989 X152 Raukumara Plain, off East Cape 36°09.70’ S 176°48.40’ E 940–820 28.11.1989 X201 Ngatoro Ridge, off East Cape 37°08.12’ S 177°17.82’ E 1050–700 06.12.1989 X207 East of Hikurangi Plateau 37°32.77’ S 176°57.70’ E 380 06.12.1989 X681 South Kermadec Ridge 35°24.91’ S 178°’39.23 E 1132–1125 13.02.1996 X768 Hikurangi Plateau 37°41.30’ S 179°00.75’ E 936 05.06.1999 Z2092 Lord Howe Seamount Chain 27°41.20’ S 155°08.80’ E 521–412 26.08.1967 Z2098 North of Three Kings Ridge 28°39.50’ S 173°01.00’ E 920 04.09.1967 Z2364 Bay of Plenty 37°25.00’ S 176°25.99’ E 277 00.00.1971 Z7187 New Zealand waters [89-BIP-1] 1989 Z7190 New Zealand waters [90-PUP] 1990 Z8487 Louisville Seamount Chain 37°40.00’ S 169°06.00’ W 1050–700 00.06.1996 Z9025 East of Three Kings Ridge 31°58.82’ S 174°15.87’ E 1680–677 00.02.1998 Z9041 Kermadec Ridge 32°10.90’ S 179°04.20’ W 260–128 05.04.1998 Z9262 Challenger Plateau, northern slope 37°29.04’ S 167°41.09’ E 904 10.09.1998 Z9263 Lord Howe Rise 34°09.20’ S 162°51.90’ E 791 18.09.1998 Z9400 Challenger Plateau, northern slope 37°18.06’ S 167°22.01’ E 755 30.09.1998 Z9742 ~3 NM east of North Cape 34°24.82’ S 173°08.00’ E 210–133 19.04.1999 Z9748 Northwest of Volkner Rocks, Bay of Plenty 37°28.01’ S 177°07.00’ E 232 30.04.1999 Z9747 ~17 NM west of Pandora Bank, Northland 34°38.91’ S 172°13.20’ E 208–198 21.04.1999 Z9751 Three Kings Islands 34°10.09’ S 172°12.04’ E 180 16.04.1999 Z9890 Western continental slope 35°00.49’ S 172°31.82’ E 167–169 30.10.1999 Z9892 Western continental slope 35°02.10’ S 172°29.25’ E 194-174 23.10.1999

97 Museum of New Zealand Te Papa Tongarewa (NMNZ) Stations

NMNZ Reg. no Geographic location Latitude Longitude Depth (m) Date NMNZ Por. 308 ~1 km off Table Cape, east of Mahia Peninsula 39°10.0’ S 178°06.0’ E 110 12.03.1957 NMNZ Por. 562 Major Island, Bay of Plenty 37°15.0’ S 176°12.0’ E 219–146 19.08.1956 NMNZ Por. 563 North of Major Island, Bay of Plenty 37°23.2’ S 176°26.15’ E 345–300 22.01.1993 NMNZ Por. 564 ~1 km off Table Cape, east of Mahia Peninsula 39°10.0’ S 178°06.0’ E 110 12.03.1957 NMNZ Por. 570 Outer Bay of Islands 35°10.0’ S 174°11.0’ E 80 16.11.1974 NMNZ Por. 571 Between Tutukaka and Poor Knights Islands 35°34.5’ S 174°39.0’ E 102 15.02.1974 NMNZ Por. 573 Ranfurly Saddle, off East Cape 37°38.0’ S 178°42.2’ E 128–126 17.01.1979 NMNZ Por. 580 Wanganella Bank, Norfolk Ridge, E slope 32°35.3’ S 167°41.8’ E 437–422 29.01.1981 NMNZ Por. 583 Middlesex Bank, NW of Three Kings Islands 34°01.4’ S 171°45.2’ E 216–201 31.01.1981 NMNZ Por. 591 Wanganella Bank, Norfolk Ridge, E slope 32°35.3’ S 167°41.8’ E 437–422 29.01.1981 NMNZ Por. 597 Wanganella Bank, Norfolk Ridge, E slope 32°41.3’ S 167°38.1’ E 296–206 30.01.1981 NMNZ Por. 615 Wanganella Bank, Norfolk Ridge, E slope 32°41.3’ S 167°38.1’ E 296–206 30.01.1981 NMNZ Por. 620 ~25 km due east of Gisborne 38°39.38’ S 178°31.62’ E 243 27.01.1995 NMNZ Por. 712 Wanganella Bank, Norfolk Ridge, E slope 32°35.3’ S 167°41.8’ E 437–422 29.01.1981

Institut de Recherche pour le Développement (IRD) LITHIST Stations

IRD Stn Geographic location Latitude Longitude Depth (m) Date CP8 Éponge Seamount, south New Caledonian slope 24°54.24’ S 168°21.35’ E 540 11.08.1999 CP9 Éponge Seamount, south New Caledonian slope 24°52.80’ S 168°21.78’ E 540–18 11.08.1999 DW5 Stylaster Seamount, south New Caledonian slope 23°38.25’ S 167°42.94’ E 433–00 10.08.1999 DW7 Éponge Seamount, south New Caledonian slope 24°55.36’ S 168°21.58’ E 530–04 11.08.1999 DW11 Kaimon Maru Seamount, south New Caledonian slop 24°46.69’ S 168°08.33’ E 254–83 11.08.1999

98 TAXONOMIC INDEX

This index has been compiled from the text. Bold numbers indicate plate or figure illustrations.

Aciculites 6, 69, 88 Gastrophanella 6, 74 ciliata 72, 74, 89 Geodia 61 higginsi 69 manawatawhi 72, 73, 74, 89, 94 Hadromerida 6 orientalis 74, 76 Herengeria 9, 29, 34, 39, 45, 88 oxytylota 74, 89 auriculata 35, 36, 41, 89, 94 papillata 74, 89 vasiformis 32, 37, 38, 39, 41, 58, 61, 89, 95 pulchra 4, 7, 41, 45, 69, 70, 71, 74, 76, 89, 95 Heterophymia 77 spinosa 74, 76 Homophymia 45, 54 sulcus 74, 75, 76, 89, 95 lamellosa 45, 47 tulearensis 74 pollubrum 89 Anaderma 56, 88 stipitata 7, 45, 46, 47, 89, 94 rancureli 89 Astrophorida 6 Isabella 88 Aulaxinia clavata 15 mirabilis 89 Awhiowhio 5, 29, 39, 45, 88 Iouea 30 osheai 32, 39, 40, 41, 89, 95 moreti 30 sepulchrum 39, 43, 44, 89, 95 Isoraphiniidae 5, 7, 61 unda 39, 41, 42, 43, 89 Azorica 77 Jereicopsis 77, 88 pfeifferae 77, 79, 80 graphidophora 89 Azoricidae 5, 76 Leiodermatium 5, 6, 76, 77, 88 Callipelta 6, 45, 47, 50, 52, 54 colini 80, 81, 82, 83, 84 cavernicola 48 crassiuscula 83 mixta 48 dampieri 77, 80, 84, 85, 89 ornata 47 intermedia 77, 78, 79, 80, 83, 84, 86 punctata 48, 49, 89, 94 linea 77, 80, 86, 89, 95 sollasi 48 lynceus 77, 79, 80 thoosa 48 pfeifferae 77, 79, 80 Corallistes 6, 7, 9, 29, 39, 88 Lepidospongia incrustans 52 australis 39, 89 Lepidothenea 52, 88 fulvodesmus 29, 30, 32 incrustans 7, 52, 53, 89, 94 multituberculatus 29, 89 Lyidium 7, 56 undulatus 29, 39, 89 torquilla 6 Corallistidae 5, 7, 29 Costifer 10, 61 Macandrewia 48, 53 vasiformis 61, 64 azorica 54 wilsoni 39, 61, 62, 63, 64, 95 spinifoliata 12, 50, 54, 55, 89, 94 Crambe 7 Mandrewiidae 5, 53 Megamorina 6 Dactylocalyx polydiscus 12 Microscleroderma 6, 65, 67, 76 Daedalopelta 45 herdmani 65, 67, 89 Discodermia 6, 7, 9, 12, 52, 53 hirsutum 65 gorgonoides 13 novaezelandiae 65, 66, 89, 95 proliferans 13, 14, 45, 89, 94 stoneae 65, 67, 89 ramifera 15 sinuosa 12 Neoaulaxinia 5, 15, 61 vermicularis 15 clavata 15, 16, 17, 19, 22, 89, 94 persicum 20, 21, 22, 27, 89, 94 zingiberadix 17, 18, 19, 22, 89, 94

99 Neopelta 50, 52, 53 Pleromidae 5, 7, 54 imperfecta 50 perfecta 50 Reidispongia 6, 27, 88 plinthosellina 50, 52, 89 coerulea 6, 27, 28, 29, 48, 84, 89, 95 pulvinus 50, 51, 89, 94 Rhizomorina 6 Neopeltidae 5, 45, 53 Neophrissospongia 30, 88 Scleritoderma 6, 67 microstylifer 30, 89 camusi 67, 89 Neoschrammeniella 6, 29, 30, 37, 45, 58, 88 flabelliformis 67, 68, 69, 89, 95 antarctica 5, 32, 33, 34, 88, 95 nodosum 67 castrum 32, 89 Scleritodermidae 5, 7, 64 fulvodesmus 30, 31, 32, 34, 72, 89, 95 Siphonidium 74 moreti 32, 39, 89 capitatum 74 norfolkii 32, 34, 89 Neosiphonia 5, 6, 22, 88 Tetracladina 6 fruticosa 24 Theonella 6, 7 motukawanui 22, 24, 25, 26, 27, 45, 89 Theonellidae 5, 12 schmidti 24, 26 superstes 6, 22, 23, 24, 27, 89, 94 Vetulina 6, 7 oamaruensis 7 Phymatellidae 5, 7, 15 stalactites 6 Pleroma 6, 7, 10, 45, 56, 88 aotea 7, 57, 59, 60, 61, 89 menoui 6, 39, 57, 58, 59, 61, 89, 95 torquilla 6 turbinatum 56, 57, 59, 61, 64, 89, 95

100